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of the 



From the collection of the 


v Jjibrary 
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San Francisco, California 


QE881 Harper 

H33 Extinct and vanishing 

mammals of the old world 







,."***? ;::; 

KORDOFAN GIRAFFE (Giraffa camelopardalis antiquorum) 

'Whenever I have watched them feeding on the tall feathery-leaved acacias, to which 
they are very partial, or stalking slowly and majestically through the park-like 
country they very commonly frequent, giraffes have always appeared to be amongst 
the most graceful and beautiful of all wild creature*." FREDERICK C. SELOUS, 1914. 

G p C f-i I- F D K G f J (E 


of the 









illustrations by 




35UOK -vIS 

FTlhere are no words that can tell the hidden 
J. spirit of the wilderness, that can reveal its 
mystery, its melancholy, and its charm. There is 
delight %i the hardy life of the open, in long rides 
, rifle in hand, in the thrill of the fight with danger- 
ous game. Apart from this, yet mingled with it, 
is the strong attraction of the silent places, of the 
large tropic moons, and the splendour of the new 
stars; where the wanderer sees the awful glory of 
sunrise and sunset in the wide waste spaces of the 
earth, unworn of man, and changed only by the 
slow changes of the ages from time everlasting. 

African Game Trails 



E1933 Dr. John C. Phillips, a founder and the first chairman 
>f the American Committee for International Wild Life Protec- 
tion, now in its twelfth year, was an official observer for our 
Government at the meetings of the London Convention for the 
Protection of the Fauna and Flora of Africa. He returned from 
that conference, which concerned itself primarily with the larger 
mammals of Africa, with the conviction that there was a basic need 
for the compilation of our present knowledge concerning the recently 
extinct and vanishing mammals, if we are to plan intelligently for 
the future preservation of wild life in this fast-changing world. 
This would be a pioneer job requiring the use of widely scattered 
sources. Such a compilation could serve as a sound foundation for 
future plans that would have to be developed to meet the ever- 
increasing threats of extermination. This research could also spot- 
light the species that are most threatened and reveal probable 
causes of extinction that might suggest new lines of effective action 
to improve their chances of survival. 

For this task the American Committee engaged the services of 
Dr. Francis Harper, an experienced mammalogist and a meticulous 
research worker. Dr. Harper started the project in May, 1936, 
and devoted more than three years to the work. The magnitude 
of the undertaking proved to be much greater than originally ex- 
pected, and the reasons for this are clearly set forth by the author 
in the introduction to the present volume. He has spoken for the 
Committee in the acknowledgments of assistance. 

The American Committee takes this opportunity to repeat its 
expression of gratitude to Dr. Harper for the hard work and care 
that he has devoted to the preparation of this volume. We are 
likewise grateful to Mr. Paul H. Oehser, editor of the United 
States National Museum, for the supervision of this volume through 
the press and for the preparation of the index. 

This whole undertaking would not have been possible without 
generous financial assistance. This has come from about 40 different 
sources, including the American Philosophical Society, the Academy 
of Natural Sciences of Philadelphia, the Boone and Crockett Club, 
the Conservation Committee of the New York Zoological Society, 
the American Wild Life Institute, and several members and mem- 


her organizations of the American Committee, as well as special 

On account of its length the publication committee decided to 
publish Extinct and Vanishing Mammals in two volumes. The 
late Dr. Glover M. Allen, in a large measure, prepared the volume 
on Extinct and Vanishing Mammals of the Western Hemisphere, 
including also certain marine mammals of all the oceans. The 
New World volume of more than 600 pages was published in 1942 
as Special Publication No. 11 of the American Committee for 
International Wild Life Protection. It was dedicated to the late 
Dr. John C. Phillips. 

The Committee appreciates the fact that ever-changing condi- 
tions require additions and supplements to the data in these volumes 
in order to bring them up to any given date. Nevertheless, keep- 
ing the information current will be a small task compared with 
the historical study, the verification of references, the biblio- 
graphical research, and the evaluation and compilation of informa- 
tion carried out by Harper and Allen in their pioneer work on the 
(recently) extinct and vanishing mammals of the Old World and 
the Western Hemisphere. 

It is our sincere hope that these volumes may serve as a founda- 
tion of information on which will be built future plans for the preser- 
vation of vanishing species of mammals in their native habitats. 
In many cases this may be most effectively brought about within a 
framework of international cooperation such as the London Con- 
vention or the Inter-American Convention. In other instances 
a threatened species may be regarded as a sort of international 
trust by the country under whose jurisdiction it may fall. For 
example, if the Great Asiatic One-horned Rhinoceros (Rhinoceros 
unicornis) should vanish from the earth (very few hundred survive 
today) it would be a world calamity and not of concern merely to 
the ruler of Assam who controls their last principal hide-out. 

International wild-life conservation should be a concern of all 
people! We must keep faith with our wild-life heritage and pre- 
serve it for the wise use of generations to come! 

(for the Committee) 
Washington, D. C. 
April 20, 1945 
Publication Committee: 






Origin, plan, methods 1 

Acknowledgments 7 

Factors in the progressive depletion of the Old World's mammalian 

faunas 8 

Australia 9 

Malay Archipelago 10 

Asia 11 

Europe 13 

Africa 15 

Madagascar 16 

The chronology of extinction 17 

The record of extinction by families 21 

Summary and conclusions 22 



Order MARSUPIALIA : Marsupials 25 

Family Dasyuridae : Dasyures, etc 25 

Genus Antechinus : Broad-footed Marsupial Mice 25 

Genus Phascogale: Brush-tailed Marsupial Rats 26 

Genus Sminthopsis : Sminthopses 29 

Genus Dasyurus : Native Cats 32 

Genus Sarcophilus : Tasmanian Devil 38 

Genus Thylacinus : Tasmanian Wolf 40 

Family Myrmecobiidae : Marsupial Anteaters 43 

Genus Myrmecobius : Numbats 43 

Family Peramelidae : Bandicoots 47 

Genus Perameles. Bandicoots 47 

Genus Macrotis: Rabbit-bandicoots or Bilbies 51 

Genus Chaeropus : Pig-footed Bandicoots 58 

Family Phalangeridae : Phalangers, etc 60 

Genus Tarsipes : Honey Possum 60 

Genus Gymnobelideus : Leadbeater's Opossum 61 

Genus Pseudocheirus : Ringtails 63 

Family Phascolarctidae : Koalas 64 

Genus Phascolarctos : Koalas 64 



Order MARSUPIALIA Continued. PAGE 

Family Vombatidae : Wombats 72 

Genus Vombatus : Common Wombats 72 

Genus Lasiorhinus : Hairy-nosed Wombats 74 

Family Macropodidae : Kangaroos, Wallabies, etc 77 

Genus Bettongia : Rat-kangaroos 77 

Genus Aepyprymnus : Rufous Rat-kangaroo 84 

Genus Potorous : Rat-kangaroos 86 

Genus Caloprymnus : Desert Rat-kangaroo 91 

Genus Lagorchestes : Hare- wallabies 93 

Genus Lagostrophus : Banded Wallaby 96 

Genus Onychogalea : Nail-tailed Wallabies 98 

Genus Petrogale : Rock-wallabies 102 

Genus Thylogale: Pademelons 106 

Genus Wallabia : Wallabies 114 

Genus Macropus : Kangaroos 120 

Order INSECTIVORA : Insectivores 122 

Family Soricidae : Shrews 122 

Genus Crocidura : Musk-shrews 122 

Order PRIMATES : Primates 123 

Family Lemuridae : Lemurs 123 

Genus Microcebus : Dwarf Lemurs 123 

Genus Cheirogaleus : Mouse Lemurs 126 

Genus Phaner : Fork-marked Lemur 129 

Genus Hapalemur: Gentle Lemurs 130 

Genus Lemur: True Lemurs 132 

Genus Lepilemur : Sportive Lemurs 144 

Family Indriidae: Sifakas, Indri, and Avahis 146 

Genus Propithecus : Sifakas 146 

Genus Indri : Indri 155 

Genus Avahi : Avahis 156 

Family Daubentoniidae : Aye-aye 158 

Genus Daubentonia : Aye-aye 158 

Family Colobidae: Leaf-eating Monkeys 160 

Genus Colobus : Colobus Monkeys 161 

Family Pongidae : Anthropoid Apes 164 

Genus Pongo : Orang-utan 164 

Genus Gorilla : Gorillas 168 

Genus Pan : Chimpanzees 176 

Order EDENTATA : Edentates 181 

Family Manidae : Pangolins 181 

Genus Smutsia: Giant and South African Pangolin 181 

Genus Phataginus: Three-cusped Pangolin 186 

Genus Uromanis : Long-tailed Pangolin 189 



Order RODENTIA : Rodents 190 

Family Leporidae : Hares and Rabbits 190 

Genus Pentalagus : Amami Hare 190 

Family Castoridae: Beavers 191 

Genus Castor: Beavers 191 

Family Cricetidae : Hamsterlike Rodents 200 

Genus Lophiomys: African Maned Rats 200 

Family Muridae: Old World Rats 205 

Genus Rattus: Typical Rats 205 

Genus Mastacomys : Broad-toothed Rats 210 

Genus Zyzomys: White-tailed Rats 211 

Order CARNIVORA : Carnivores 211 

Family Canidae : Wolves and Foxes 211 

Genus Simenia : Simenian Foxes 212 

Genus Canis: Wolves 213 

Genus Nyctereutes : Raccoon-dogs 215 

Family Ursidae : Bears 217 

Genus Ursus : Bears 217 

Family Mustelidae : Weasels, etc 232 

Genus Mustek: Weasels, Minks, and Stoats 233 

Genus Martes : Sables and Martens 235 

Genus Gulo : Wolverines 241 

Family Viverridae: Civets, Mongooses, etc 244 

Genus Arctictis : Binturongs 244 

Genus Fossa : Fossane 249 

Family Protelidae : Aard- wolves 250 

Genus Proteles : Aard- wolves 250 

Family Felidae: Cats 254 

Genus Cryptoprocta : Fossa 254 

Genus Felis: Cats 256 

Genus Lynx : Lynxes 265 

Genus Caracal : Caracals 272 

Genus Acinonyx : Cheetahs 274 

Genus Leo : Lions 288 

Genus Panthera : Leopards and Tigers 299 

Order PROBOSCIDEA : Proboscideans 310 

Family Elephantidae : Elephants 310 

Genus Elephas: Asiatic Elephants 311 

Genus Loxodonta: African Elephants 316 

Order PERISSODACTYLA : Odd-toed Ungulates 322 

Family Equidae: Horses, Zebras, and Asses 322 

Genus Equus: Horses, Zebras, and Quagga. . . : 322 

Genus Asinus: Asses . . 345 



Family Tapiridae : Tapirs 371 

Genus Acrocodia: Malay Tapir 372 

Family Rhinocerotidae : Rhinoceroses 375 

Genus Rhinoceros: Asiatic One-horned Rhinoceroses 375 

Genus Dicerorhinus : Asiatic Two-homed Rhinoceroses 390 

Genus Diceros : Black Rhinoceroses 396 

Genus Ceratotherium : White Rhinoceroses 402 

Order ARTIODACTYLA : Even-toed Ungulates 414 

Family Hippopotamidae : Hippopotamuses 414 

Genus Hippopotamus: Common Hippopotamuses 414 

Genus Choeropsis: Pygmy Hippopotamus 419 

Family Camelidae : Camels and Llamas 421 

Genus Camelus : Camels 421 

Family Tragulidae : Chevrotains 425 

Genus Hyemoschus : Water Chevrotains 425 

Family Moschidae : Musk Deer 427 

Genus Moschus: Musk Deer 427 

Family Cervidae : Deer 435 

Genus Muntiacus: Muntjaks 435 

Genus Rucervus : Swamp Deer 436 

Genus Cervus : Red Deer and Sikas 443 

Genus Elaphurus : Pere David's Deer 467 

Genus Rangifer : Reindeer 469 

Family Giraffidae : Giraffes and Okapi 484 

Genus Giraffa : Giraffes 484 

Genus Okapia : Okapi 506 

Family Bovidae: Cattle, Sheep, Goats, and Antelopes 510 

Genus Novibos: Cambodian Wild Ox 510 

Genus Bos: Cattle 511 

Genus Bibos: Gaurs and Bantengs 514 

Genus Poephagus : Yaks 528 

Genus Bison : Bisons 531 

Genus Bubalus : Asiatic Buffaloes 538 

Genus Anoa : Dwarf Buffaloes 548 

Genus Syncerus : African Buffaloes 554 

Genus Ovis : Sheep 557 

Genus Ammotragus : Audads 600 

Genus Capra : Goats and Ibexes 606 

Genus Capricornis : Serows 635 

Genus Alcelaphus : Hartebeests 642 

Genus Damaliscus : Bontebok and allies 653 

Genus Connochaetes : Gnus 659 

Genus Cephalophus : Duikers 663 


Order ARTIODACTYLA Continued. 

Family Bovidae Continued. PAGE 

Genus Oreotragus : Klipspringers 668 

Genus Nesotragus : Sunis 672 

Genus Dorcatragus : Beira 674 

Genus Ammodorcas : Dibatag 675 

Genus Saiga : Saiga Antelope 677 

Genus Gazella: Gazelles 683 

Genus Aegoryx: White Oryx 690 

Genus Oryx: Oryxes 693 

Genus Hippotragus : Roan and Sable Antelopes and Blaauw- 

bok 698 

Genus Addax : Addax 711 

Genus Tragelaphus : Bushbucks 716 

Genus Taurotragus : Elands 722 


INDEX . 811 



26. Binturong: Arctictis binturong subsp. (After photograph in 

Brehm and specimen in Philadelphia Zoo) 246 

27. Aard-wolf: Proteles cristatus subsp 253 

28. Fossa: Cryptoprocta ferox Bennett (After photograph in Brehm) .255 

29. European Lynx : Lynx lynx lynx (Linnaeus) (After Standard Nat. 

Hist., etc.) 266 

30. Barbary Lynx: Caracal caracal algirus (Wagner) (From specimen 

in Philadelphia Zoo) 273 

31. Indian Cheetah: Acinonyx jubatus venaticus (Hamilton Smith). 284 

32. King Cheetah: Acinonyx rex Pocock (After Pocock, 1927) 287 

33. Mongolian Wild Horse: Equus przewalskii Poliakov (After photo- 

graph in Brehm) 326 

34. Quagga: Equus quagga Gmelin (After Standard Nat. Hist.) 335 

35. Burchell's Zebra: Equus burchellii burchellii (J. E. Gray) (After 

Brehm) 340 

36. Mountain Zebra: Equus zebra zebra Linnaeus (After photo- 

graphs in Brehm and Newnes) 343 

37. Nubian Wild Ass: Asinus asinus africanus Fitzinger 346 

38. Indian Wild Ass: Asinus hemionus khur (Lesson) 365 

39. Malay Tapir: Acrocodia indica (Desmarest) 373 

40. Great Indian Rhinoceros : Rhinoceros unicornis Linnaeus 376 

41. Javan Rhinoceros: Rhinoceros sondaicus Desmarest 382 

42. Northern White Rhinoceros: Ceratotherium simum cottoni 

(Lydekker) (After Lang) 408 

43. Pygmy Hippopotamus: Choeropsis liberiensis (Morton) 420 

44. Water Chevrotain: Hyemoschus aquaticus subsp. (After Brehm) . 426 

45. Schomburgk's Deer: Rucervus schomburgki Blyth 437 

46. Barbary Stag : Cervus elaphus barbarus Bennett 458 

47. Pere David's Deer: Elaphurus davidianus Milne Edwards 468 

48. Spitsbergen Reindeer: Rangifer platyrhynchus (Vrolik) (After 

Wollebaek, 1926) 480 

49. Southern Giraffe: Giraffa camelopardalis capensis (Lesson) (After 

Brehm, Lydekker, etc.) 504 

50. Okapi: Okapia johnstoni (P. L. Sclater) 507 

51. Caucasian Bison: Bison bonasus caucasicus Hilzheimer 538 

52. Common Anoa: Anoa depressicornis (Hamilton Smith) 551 

53. European Mouflon: Ovis musimon (Pallas) 575 

54. Barbary Sheep : Ammotragus lervia subsp 600 

55. Mediterranean Ibex: Capra pyrenaica hispanica Schimper (After 

Lydekker and Ward) '. 613 

56. Abyssinian Ibex: Capra walie Riippell (From photograph, Field 

Museum) * 624 

57. Markhor: Capra falconeri subsp. (After Cassell) 628 


58. Sumatran Serow: Capricomis sumatraensis sumatraensis (Bech- 

stein) (After Mohr, 1934) 636 

59. Bubal Hartebeest: Alcelaphus buselaphus buselaphus (Pallas) 

(After photograph in Brehm) 644 

60. Bontebok: Damaliscus dorcas (Pallas) (After Brehm and 

Lydekker) 655 

61. White- tailed Gnu: Connochaetes gnou (Zimmermann) 662 

62. Saiga: Saiga tatarica (Linnaeus) (After Brehm) 681 

63. White Oryx: Aegoryx algazel (Oken) (From specimen in Phila- 

delphia Zoo) 692 

64. Blaauwbok: Hippotragus leucophaeus (Pallas) (After Daniell, in 

Jardine's Naturalist's Library) 699 

65. Giant Sable Antelope: Hippotragus variani Thomas 709 

66. Addax: Addax nasomacidatus (Blainville) (After Brehm, etc.) .. 713 

67. Nyala: Tragelaphus angasii Angas (After Sclater and Thomas, 

1900) 718 




THE present work had its origin in a strongly felt need for defi- 
nite information on the mammals that have become extinct 
during the Christian Era, on those that are now threatened with the 
same fate, on the factors contributing to the progressive depletion of 
the world's mammalian faunas, and on the measures that have been 
hitherto or may be hereafter undertaken for their preservation. 
It consists to a large extent of an inventory of vanishing resources, 
as an essential step in their conservation. 

The plan and the inception of this investigation are due to the 
keen interest and foresight of the late Dr. John C. Phillips, founder 
and first chairman of the American Committee for International 
Wild Life Protection. The work has been carried out under the 
auspices of that organization and has been supported in part by 
a grant from the Penrose Fund of the American Philosophical 
Society. The Academy of Natural Sciences of Philadelphia was 
chosen as the headquarters of the investigation, largely on account 
of the very exceptional resources of its library in the literature of 
natural history. 

As originally projected, the investigation was to have covered the 
entire world and the results were to be published in a single volume. 
Owing to limitations of time, space, and available funds, as well 
as the unforeseen magnitude of the task, the present volume is 
restricted to the mammals of the Old World. The major part of 
my work was concluded early in 1939; in only a few instances, 
therefore, has it been possible to take into account the subsequently 
published literature. Another volume, prepared in large part by 
Dr. Glover M. Allen, late curator of mammals at the Museum of 
Comparative Zoology, and published in December, 1942, deals with 
the mammals of the New World and with the marine forms. 

It was also hoped to include in the Introduction a general sur- 
vey of conservation conditions so far as they affect mammals 
in the various countries of the world. Although it has not been 
possible to carry out this feature, fortunately the need for it has 



been obviated in part by Brouwer's The Organisation of Nature 
Protection in the Various Countries (1938) - 1 

In the preparation of the accounts of the various mammals 
treated, the aim has been to assemble and to present in concise form 
such information as could be obtained on the following points: 

Former range and numbers; 

Present range and numbers (of vanishing species) ; 

Date and rate of disappearance in each country (of species that have 

become extinct, either locally or completely) ; 
Causes of depletion or extinction, either direct or indirect; 
Economic uses or importance; 
Esthetic considerations; 
The meiBures that have been or might be undertaken for the preservation 

of each vanishing species. 

The primary source of this information has been the published 
literature. For this purpose, the library of the Academy of Natural 
Sciences of Philadelphia has been the mainstay. In addition, I have 
drawn to some extent upon the library resources of the United States 
National Museum, the Museum of Comparative Zoology, the Ameri- 
can Museum of Natural History, the Charles Sheldon Collection 
at Yale University, and the American Philosophical Society. 

An especially valuable source of information has been corres- 
pondence with zoologists and conservation officials in most of the 
countries of the Old World. By means of questionnaires, distributed 
for the most part through the collaboration of the International 
Office for the Protection of Nature in Brussels, a great mass of 
fresh and largely unpublished data on the distribution, numbers, 
economic status, and conservation of mammals has been assembled. 

The unselfish cooperation of these contributors, on a scale per- 
haps unprecedented in this field, has been an extremely helpful 
and highly appreciated feature of the investigation. 

Additional material and documents bearing upon the present 
subject had been accumulating for some years in the office of the 
American Committee for International Wild Life Protection, and 
these have been utilized to considerable advantage. 2 

Perhaps few zoologists have had better occasion than myself to 
become impressed with the inexhaustible nature (and at the same 
time the inadequacy) of the literature on systematics, distribution, 

1 Special publication of the American Committee for International Wild Life 
Protection, No. 9. 

2 Dr. Glover M. Allen, in making use of office data of this sort in his com- 
panion volume on mammals of the Western Hemisphere (1942), seems to have 
been under the erroneous impression that I was responsible for gathering prac- 
tically all of them, and consequently he has mentioned my name with the best 
of intentions but with considerably greater frequency than the facts would 
warrant. Credit for many of the data is due to sources indicated above. 


economics, life histories, and related phases in the study of mammals. 
Likewise few can become more conscious than myself of the incom- 
pleteness of the present report on the points it endeavors to cover. 
The chief handicap has been the sheer limitations of time, despite 
unremitting labor during a period of practically three years. Sec- 
ondary handicaps have been the nonavailability of certain litera- 
ture, and the virtually unusable nature (to an Anglo-Saxon) of 
much of the literature in the Slavic and Oriental languages. 

In nearly every case a separate account has been provided for 
each species or subspecies coming within the scope of the present 
report. In matters of taxonomy and nomenclature I have endeav- 
ored to follow the best authorities available, as exemplified in recent 
monographs, catalogues, or check-lists. However, unanimity of 
opinion on every detail is not to be expected of the specialists in this 

A really surprising amount of confusion in the nomenclature of 
even some of the largest and best-known of the Old World mammals 
has come to light as an incidental feature of the present investi- 
gation. This seems to be due largely to lack of proper attention to 
type descriptions and type localities, and to some extent to dis- 
regard of the International Rules of Zoological Nomenclature. I 
have attempted to straighten out some of the major nomenclatural 
difficulties in two preliminary papers (Harper, 1939, 1940) , while a 
few minor points, relating especially to type localities, are touched 
upon in the present work. 

Each account furnishes, in addition to the technical name of the 
mammal under discussion, its common names in English and (if 
known) in French, German, Italian, and occasionally other lan- 
guages of western Europe. No attempt has been made to compile 
names from unfamiliar or unwritten languages, and in only a few 
exceptional cases have any been included. This statement, however, 
does not apply to such native names as may have been taken over 
bodily into the English or other European languages. 

After the common names comes the original reference, or citation 
of the type description. I have been able to verify probably 95 
percent of these original references in the library of the Academy 
of Natural Sciences of Philadelphia, and a few others elsewhere. 
A statement of the type locality is then added in parentheses ; as far 
as feasible, it is given in the form of an exact quotation from the 
original description. In many cases brief supplementary or explana- 
tory remarks are called for. 

No attempt has been made to supply a complete list of synonyms, 
and usually none whatever are cited. In certain cases, however, 
where circumstances seem to render it advisable, one or more 
synonyms are cited. For example, if the name considered valid and 


adopted here happens to be less familiar than one or more that are 
replaced, the latter may be cited. Or, if certain recently proposed 
names are not considered valid, these may likewise be cited in 

Persons interested in the study of mammals have frequent need 
of consulting good illustrations of the various species and subspecies. 
For this reason a special point has been made of supplying refer- 
ences to such illustrations. While such a list of references can rarely 
be exhaustive, it is believed that the hundreds of references given 
here include at least a majority of the good published figures or 
plates of the mammals under discussion. In the case of slightly 
differentiated subspecies, an outstanding difficulty very frequently 
encountered has been the determination of the particular one 
figured. Even if the author or artist has provided a trinomial 
designation and this is far from being a universal practice there 
is always a possibility of misidentification, or of eventual refinement 
of classification that will throw the identification into doubt, 
unless the geographical provenance of the specimen figured is 
accurately stated. The frequent disregard of this rule is the cause 
of endless vexation, and it detracts seriously from the value of the 
figures as zoological illustrations. Thus, in many cases the sub- 
specific identity of a figure mentioned in the references is in doubt, 
and some of these cases are indicated by a parenthetical query: 
"subsp.?" This uncertainty of identification extends inevitably 
even to some of Mr. Poole's excellent drawings that illustrate the 
present text. For some are based upon "zoo" specimens, many of 
which are notoriously of uncertain provenance or even represent the 
hybrid offspring of different subspecies in captivity ; while others are 
based upon previously published figures, themselves of somewhat 
uncertain subspecific identity. 

A brief description of each species or subspecies is included. In 
its preparation I have aimed to utilize the type description so far 
as it is at all adequate; but in many cases later and more complete 
or more accurate descriptions have necessarily been drawn upon 
for at least some of the characters. Constant caution is required, 
however, in making use of reviews, catalogues, or monographs in 
which the descriptions may be based upon specimens of unspecified 
provenance. In all possible cases I have indicated the source of the 
information by a direct quotation or, in the case of translation or 
paraphrasing, by at least a bibliographical reference. 

It may be remarked here that the entire report is documented 
with such references to the fullest possible extent, not only as a 
matter of simple justice to the authors of the works drawn upon, 
but as an essential aid to the reader in verifying statements, and in 
ascertaining what source material has been utilized on the one hand, 


or overlooked or disregarded on the other hand. The common 
literary sins of failing to acknowledge sources of information, of 
giving incomplete references, and of taking liberties with quotations, 
have been scrupulously avoided as far as has lain within my power. 
These matters have called for the closest possible attention in a 
work that is so largely a compilation as the present one. 

Perhaps no two mammalogists would agree completely on just 
what species or subspecies come properly within the scope of this 
report. In the first place, that scope is not completely explained in 
the rather brief title chosen. With the exception of a few partially 
aquatic species, such as the hippopotamuses, only land mammals are 
included. The various marine and fresh-water species are dealt with 
in Dr. Allen's volume (1942). A somewhat more exact but unduly 
awkward title might have been Land Mammals of the Old World 
that are Extinct, or Vanishing, or in Need of Special Protection. 
Some of the forms included are no doubt actually increasing under 
protection at the present moment but nevertheless deserve and 
require the fullest possible care in order that they may continue to 

It has been deemed advisable to include all African mammals 
accorded protection in Schedules A and B of the London Convention 
of 1933, even if subsequent investigation has shown that certain 
forms are in less urgent need of close protection than was at first 
supposed. On the o^her hand, the simple limitations of time and 
funds have excluded a certain number of rare and more or less en- 
dangered species whose status is probably more unsatisfactory than 
that of a good many included species. 

Finally, there are doubtless a considerable number of other mam- 
mals (especially small, inconspicuous, or secretive species) that have 
progressed far toward the vanishing point, or that have actually 
become extinct, without their status having become known to zoolo- 
gists. There is no royal road to the discovery of such a state of 
affairs. Time and again extinction has taken place years in advance 
of the fact coming to scientific attention. Thus, at the very best, 
the present report could embody no more than a certain portion of 
the current (and decidedly incomplete) knowledge on the subject. 

A few words may be said here on the difficult subject of the 
arrangement or sequence of the systematic groups families, genera, 
species, and subspecies. The present arrangement of families is ac- 
cording to Simpson (1931). Beyond this point there is apparently 
no single, comprehensive, up-to-date guide to be followed. Many 
recent authors of faunal lists or catalogues do not even undertake 
an explanation of the sequence they adopt. For the large group of 
ungulate mammals Lydekker's well-known catalogue (1913-1916) 
furnishes a convenient guide in the arrangement of genera, species. 


and subspecies. In the same way Iredale and Troughton's Australian 
check-list (1934) serves for the marsupial groups. In other groups 
(such as the Carnivora and the Primates) I have merely attempted 
to follow general usage in so far as any such usage has been 
discoverable. In the category of subspecies, the original, "nominate," 
or "typical" subspecies is introduced first, and is followed by the 
others, usually in a more or less geographical sequence from north to 
south or east to west. 

Under each family heading a brief paragraph has been introduced, 
stating the general distribution of the family, the number of genera 
and species or subspecies it contains (exclusive of fossil forms) , and 
the number of forms that have called for discussion in Dr. Allen's 
preceding volume and in the present volume. The number so 
treated varies from one in each of several families to more than a 
hundred in the cattle family (Bovidae). Since no indigenous land 
mammals occur in Antarctica, there is no need of further mention 
of this region in the distributional statements. 

The 1933 London Convention for the Protection of the Fauna and 
Flora of Africa is mentioned with considerable frequency in this 
volume. Since some readers may not be familiar with this Con- 
vention and its far-reaching importance in the cause of international 
wildlife preservation, a few words of explanation are inserted here. 
The conference was called by invitation of Great Britain and was 
attended by accredited representatives of the nine countries having 
territories in Africa. The Convention became effective in January, 
1936, when it had been ratified by five of the nine participating 
governments. By January, 1940, ratification by three more countries 
had taken place. Among the measures agreed upon by the Con- 
vention are the establishment of national parks and nature reserves, 
the regulation of traffic in animals, and the prohibition of encircling 
fires and (wherever possible) of the use of poison, dazzling lights, 
nets, and traps for hunting animals. The Annex to this Convention 
lists, as Class A species, 17 mammals, 3 birds, and 1 plant, for 
which rigid protection is agreed upon. It also lists, as Class B 
species, 13 mammals and 9 birds which, although not requiring 
such rigid protection, may be taken only under a special form of 
license. The number of mammals so protected is actually much 
larger than indicated in the above statements; for example, the 
40 forms of Madagascar lemurs count as only a single item in the 
list, and the same is true of the dozen subspecies of Giraffes. Further 
details are set forth in Special Publications 6 and 10 of the American 
Committee for International Wild Life Protection (1935 and 1940). 



The chief burden of promoting the present investigation, in both 
a spiritual and a financial sense, was magnanimously assumed in 
the place by the late Dr. John C. Phillips, not merely in his 
capacity as chairman of the American Committee for International 
Wild Life Protection, but as a more or less personal responsibility. 
The main lines of the investigation have been carried out as origi- 
nally planned by him. Other members of the Committee have also 
made generous contributions of funds, information, and advice. 
When the magnitude of the task began to exceed all original esti- 
mates, a grant from the Penrose Fund of the American Philosophical 
Society provided timely aid. After Dr. Phillips's death in November, 
1938, a subcommittee, consisting of Dr. Alexander Wetmore, Mr. 
Charles M. B. Cadwalader, and Mr. Harold J. Coolidge, Jr., by 
vigorous action found the means for completing the investigation. 

I am further and particularly indebted to Mr. Cadwalader, as 
director and president of the Academy of Natural Sciences of Phila- 
delphia, for the provision of desk space and library facilities in 
this institution. The already great resources of the Academy's 
library have been considerably augmented, during and in behalf of 
this investigation, by the acquisition of numerous important works 
on mammals, through the efforts of Mr. Cadwalader, Mr. Brooke 
Dolan, II, and Mr. George L. Harrison. 

The whole-hearted cooperation of the International Office for the 
Protection of Nature in Brussels, and particularly of its Secretary, 
Mrs. Tordis Graim, is most gladly and gratefully acknowledged. 
Mrs. Graim has generously undertaken and admirably fulfilled the 
task not only of distributing questionnaires to numerous zoologists 
and conservation officials in the Old World, but also of translating 
and compiling the very valuable data thus obtained. 

Through the courtesy of Dr. H. E. Anthony and Mr. George G. 
Goodwin, of the American Museum of Natural History, lengthy 
portions of indispensable works in Russian by Ognev and Nasonov 
have been translated at that institution and placed in my hands. 
Thereby a great deal of important information, not generally 
available to non-Russian zoologists, has been incorporated in the 
pages of the present work. 

I must not omit to mention the patience and accommodation of 
Dr. Remington Kellogg, of the United States National Museum, 
during the hours I have spent in his office, consulting various works 
not available in Philadelphia. 

No words of mine can add to the value of the drawings produced 
by the masterful strokes of Earl Poole's pen. They will be appre- 
ciated by the reader not only as unusually faithful delineations of 


the mammals they represent, but as a welcome embellishment of 
the long pages of text. 

The host of correspondents and collaborators, who have contrib- 
uted first-hand and hitherto unpublished information of very ex- 
ceptional value, and whose names will be found in proper place 
on scores of the following pages, deserve the highest gratitude of 
the sponsors and the compiler of this report. Without their contri- 
butions the work would have been deprived of one of its most 
essential features. 

Finally, the most cordial thanks are due to the various authors 
and publishers whose books and papers have been utilized in the 
preparation of this work. It is hoped that they will be rewarded in 
part, at least, by the complete acknowledgment of all items of 
information so derived. 

Of the following accounts of Old World species, 17, which 
Dr. Francis Harper did not have opportunity to prepare on account 
of taking up other investigations, were written by Glover Morrill 
Allen and are subscribed with his initials. These accounts are in 
large part based on the data already brought together by Dr. Harper, 
to whom every credit is due for the extensive research and corres- 
pondence which he undertook in order to assemble the essential 
facts. The 17 accounts are: Crocidura juliginosa trichura, Christ- 
mas Island Shrew; Rattus macleari, Captain Maclear's Rat; Rattus 
nativitatis, Christmas Island Burrowing Rat; Colobus polykomos 
and Colobus badius races, Colobus Monkeys; Pan troglodytes and 
races, the Chimpanzee; Pongo pygmaeus, the Orang-utan; Hippo- 
potamus amphibius and races, the Hippopotamuses; Choeropsis 
liberiensis, Pygmy Hippopotamus ; Hyemoschus aquations and races, 
Water Chevrotains; Cervus elaphus barbarus, North African Red 
Deer; Loxodonta africana africana, South African Bush Elephant; 
Diceros bicornis and races, Black Rhino ; Equus burchellii burchellii, 
Burchell's Zebra; Equus zebra and race, Mountain Zebra; Equus 
quagga, the Quagga; Oryx gazella and race, Gemsbok; Aegoryx 
algazel, Scimitar Oryx; and Syncerus caffer caffer, Cape Buffalo. 

G. M. A. 


IN the course of the present studies on the mammals that have 
become extinct during the Christian Era, and on others that are 
now threatened with the same fate, it has become convincingly 
evident that the process of extinction is taking place at a steadily 
accelerated rate. During this period of approximately 2,000 years, 


the world has lost, through extinction, about 106 known forms of 
mammals. About 28 percent of these are subspecies of still existing 
species, but the full species completely and irretrievably lost number 
approximately 77. 

Between A. D. 1 and 1800, about 33 mammals are more or less 
definitely known to have become extinct (see list, pp. 17-18) . Each 
half -century period since 1800 shows a steadily increasing rate of 
extinction. The last 100 years have witnessed the passing of about 
67 percent of the 106 extinct forms. In the past 50 years approxi- 
mately 38 percent as many forms have been exterminated as in all 
previous recorded history. At the present time more than 600 others 
require consideration as vanishing or threatened forms. It is well 
within the bounds of possibility that during the next hundred years 
we may be extinguishing this group at the approximate rate of one 
form per year. 

In seeking the causes of this world-wide tragedy, it becomes ap- 
parent that conditions vary widely over the different regions of 
the globe, although there is a single major underlying factor nearly 

For the purposes of the present inquiry, we may here pass briefly 
in review the major regions that are covered in this volume: 
Australia, the Malay Archipelago, Asia, Europe, Africa, and Mada- 


Conditions in Australia are peculiar and exceptional, owing to 
the fact that its unique native mammalian fauna is predominantly 
marsupial, and so lowly organized as to be quite unfitted for coping 
with certain exotic and aggressive species introduced by civilized 
man. The chief of these are the European Red Fox, the Domestic 
Cat, the European Rabbit, the House Rats, and the House Mouse. 
Further competition results from the encroachment of hosts of 
sheep and cattle upon the ancestral grazing grounds of the her- 
bivorous marsupials. An apparently minor predatory role is played 
by the Dingo (Cams dingo), which was presumably introduced by 
aboriginal man. 

The Fox and the Cat (which has become feral in large numbers) 
have long been active in the direct extermination of the smaller and 
comparatively helpless marsupials. The Rabbit, in millions, operates 
indirectly but no less effectively by overrunning the land, occu- 
pying all available burrows, and depriving the herbivorous mar- 
supials (even such large species as the kangaroos) of the food 
necessary to their existence. The introduced rats and mice usurp 
the habitats of the native species. Even sanctuaries are not proof 
against such enemies as the foregoing. 


The serious depletion of the native fauna by these agencies is 
supplemented by widespread bush fires, by conversion of a vast 
acreage of wild land into crop or grazing lands, by the huge fur 
trade, by epizootic disease, and by the large-scale use of poisoned 
bait, which takes toll of many animals besides the pests against 
which it is directed. 

Altogether, the situation in Australia has gotten largely beyond 
human control. The rapidly growing list of extinct forms already 
contains at least the following 11 : 

Freckled Marsupial Mouse (Antechinus apicalis) 

New South Wales Barred Bandicoot (Pemmeles jasciata) 

Western Barred Bandicoot (Perameles myosura myosura) 

Nalpa Bilby (Macrotis lagotis grandis) 

Leadbeater's Opossum (Gymnobelideus leadbeateri) 

Gaimard's Rat-kangaroo (Bettongia gaimardi) 

Gilbert's Rat-kangaroo (Potorous gilbertii) 

Broad-faced Rat-kangaroo (Potorous platyops) 

Parma Wallaby (Thylogale parma) 

Toolach Wallaby (Wallabia greyi) * 

White-tailed Rat (Zyzomys argurus argurus) 

Dr. W. K. Gregory, of the American Museum of Natural History, 
says (1924, p. 11) : "Late in the eighteenth century, there arrived 
in Australia by far the most destructive placental mammal the 
world has ever seen, Homo sapiens, variety europaeus, who has 
devastated the continent and is now completing the work of 


Insular faunas are of extraordinary interest because of their 
tendency toward endemism and because of the light they throw 
upon geological history and evolutionary processes. At the same 
time, by reason of the more or less strictly circumscribed nature of 
their habitats, and by reason of a certain lack of adaptability or 
self-defense, they are peculiarly vulnerable to attack and extermi- 
nation by enemies of foreign origin. Thus the Malay Archipelago 
commands the attention of the conservationist as well as of the 
evolutionist. Incidentally, it was in this environment, in the fertile 
mind of Alfred Russel Wallace, that one of the germs of the evolu- 
tionary idea developed. 

So far this region, containing the richest insular faunas of the 
entire world, has fared moderately or at least comparatively well, 
having lost only three mammals, all from tiny Christmas Island, 
lying some 200 miles off the south coast of Java. These are a shrew 
(Crocidura fuliginosa trichura) and two species of indigenous rats 
(Rattus macleari and R, nativitatis) , all of which have succumbed 

i A single captive remained alive in 1938 (Troughton, 1938, p. 407). 


to an invasion of House Rats and Domestic Cats, either through 
direct attack or through some epizootic introduced by one or both 
of these animals. 

On the other hand, through the archipelago generally, cultivated 
areas and the native population show a strong tendency to increase ; 
this is especially true of the Sunda Islands and the Philippines. 
Thus the native mammals are engaged in a steady retreat into the 
dwindling forests. 

In the Netherlands Indies many good protective measures have 
been adopted. No less than 76 nature reserves have been created, 
and these may be regarded as the final refuge of the native fauna. 
Hunting and export of wild animals are prohibited except under 
special license. 

In Borneo and New Guinea the native population is less dense 
than in the Sunda Islands, and there is apparently little use by the 
natives of firearms that primary factor in the extermination of 
wild life. 

The vanishing mammals of the archipelago, for which special 
concern is felt, include the following: 

Orang-utan (Pongo pygmaeus) 

Sumatran Elephant (Elephas maximus sumatranus) 

Javan Rhinoceros (Rhinoceros sondaicus} 

Sumatran Rhinoceros (Dicerorhinus sumatrensis sumatrensis) 

Babirussa (Babirussa babyrussa) 

Javan Banteng (Bibos sondaicus sondaicus) 

Bornean Banteng (Bibos sondaicus lowi) 

Tamarao or Dwarf Buffalo of Mindoro (Anoa mindorensis) 

Common Anoa of Celebes (Anoa depressicornis) 

Mountain Anoa of Celebes (Anoa jergusoni) 

Sumatran Serow (Capricornis sumatraensis sumatraensis) 

Of these, the Javan Rhin'oceros is in the most serious condition, 
being reduced to perhaps two dozen individuals. 


The fauna of this greatest of the continents has been safeguarded 
in part by natural conditions. Chief among these is the sparsity of 
the human population over such vast areas as the taiga and the 
tundra of Siberia and the deserts of Mongolia, Chinese Turkestan, 
Persia, and Arabia. The great mountain masses of the Himalaya, 
Tian Shan, and Altai systems, as well as numerous lesser ranges, 
have also afforded a measure of protection to the mammals adapted 
to these high altitudes. 

A factor in the preservation of the large mammals of Afghan- 
istan and Tibet has been the exclusion of all but a handful of 
foreigners. India, despite its teeming population, has not exter- 


minated a single mammal, thanks to the protective attitude toward 
game assumed both by the native rulers and by the British adminis- 
tration. In China, unfortunately, there seems to be little or no 
thought of the conservation of wild life on the part of the great mass 
of the population. 

One of the most decisive factors in the accelerated depletion of 
the game resources of Asia (and of other continents likewise) 
during recent years has been the increasing use of modern rifles of 
high power and precision. This has been especially noticeable in 
Tibet, according to reports of recent explorers, and also in Arabia. 
In the deserts of Iraq and Arabia pursuit of gazelles and other 
animals by motor car has recently become a very serious menace to 
their survival. 

The Asiatic rhinoceroses, the Saiga Antelope, such large horned 
ruminants as the Wapiti and other members of the deer family, and 
even the lowly pangolins, have been victimized in a peculiarly 
distressing way, merely because of the apparently wholly mythical 
value of the horns, scales, and other parts of the body in the Chinese 
pharmaceutical trade. This belief is so deeply rooted that probably 
no educational campaign would be effective in staving off the 
extermination of any species at the mercy of the peoples who regard 
powdered rhino horn, for example, as a panacea. Even in countries 
far beyond China's borders, protection of rhinoceroses and other 
species in similar demand is made extraordinarily difficult by the 
fabulous prices set upon them and by the incentive for poaching 
under these circumstances. When the last Asiatic rhino is gone, 
and the fancied benefits from its powdered horn are no longer 
available, possibly then the tragic fallacy of the whole business 
will dawn upon those responsible for the extermination of this 
section of the world's fauna. 

Of fur-bearing animals, probably the highly prized Siberian 
Sables have been subjected to severest pressure, but the Soviet 
Government has created several great reserves for their protection, 
and has maintained a closed season on Sables over the whole terri- 
tory of the USSR. 

Despite the many-sided attack upon Asiatic mammals for the 
sake of their meat, hides, fur, horns, scales, and even raw body 
fluids that continent has exterminated to date, as far as known, 
only three forms: the Japanese Wolf (Canis hodophilax), the 
Syrian Wild Ass (Asinus hemionus hemippus) , and Schomburgk's 
Deer (Rucervus schomburgki) . 

There are a number of others, however, for which the same fate 
is more or less imminent. Notable among these are the following: 

Indian Cheetah (Acinonyx jubatus venaticus) 
Asiatic Lion (Leo leo persicus) 


Przewalski's Horse (Equus przewalskii) 

Transcaspian Wild Ass (Asinus hemionus finschi) 

Indian Wild Ass (Asinus hemionus khur) 

Javan Rhinoceros (Rhinoceros sondaicus) 

Asiatic Two-horned Rhinoceroses (Dicerorhinus sumatrensis lasiotis and 

D. s. niger) 

Yarkand Stag (Cervus yarkandensis) 
McNeill's Deer (Cervus macneilli) 
White-lipped Deer (Cervus albirostris) 
Malayan Gaur (Bibos gaurus hubbacki) 
Gobi Argali (Ovis ammon darwini) 
Semipalatinsk Argali (Ovis ammon collium) 
Anadyr Bighorn (Ovis nivicola subsp.) 

There are doubtless additional forms of Asiatic Wild Sheep whose 
existence is seriously threatened, but information on the present 
status of certain ones is scarcely sufficient to warrant a definite 


In view of the fact that the European type of culture has 
generally had such a devastating effect upon native faunas wherever 
it has spread in colonies and settlements throughout the rest of the 
world, it is gratifying to find that the mammalian fauna of Europe 
itself has retrograded no further than it has. The chief impover- 
ishment has naturally occurred in the British Isles and other densely 
populated countries of Western Europe. And yet fewer Recent 
mammals have been exterminated in Europe than in North America 
or Australia or Africa. They seem to number only six, as follows: 

European Lion (Leo leo subsp.) 

European Wild Horse (Equus caballus subsp.) 

Aurochs (Bos primigenius) 

Caucasian Bison (Bison bonasus caucasicus) 

Pyrenean Ibex (Capra pyrenaica pyrenaica) 

Portuguese Ibex (Capra pyrenaica lusitanica) 

The retrogression of the European fauna has no doubt been due 
in the first place to the widespread clearing of forests and their 
replacement by lands devoted to habitations, transportation systems, 
crops, or grazing. Hunting, however, has constituted the most im- 
portant part of the direct human pressure upon the wild animals. 
While this sort of pressure began to be felt ages ago, it was primarily 
the invention and improvement of firearms that enabled man to 
proceed with ever-increasing rapidity on his course of extermination. 
Species of comparatively large size, furnishing valuable meat and 
hides, have been the prinicipal sufferers. Thus four of the six 
extinct European mammals are members of the cattle family 


In Europe, as contrasted with the United States, there is a far 
greater proportion of closely guarded private estates, and hunting 
of large game is chiefly restricted to the wealthy few. This con- 
dition of affairs has*resulted in a much slower rate of extermination 
than in the United States, despite the large number of national 
parks and wild-life refuges in this country. Furthermore, the 
European attitude appears much more tolerant toward such preda- 
tory animals as Wolves and Brown Bears, which have been able 
to survive so far in such countries as Spain, France, Italy, Yugo- 
slavia, Greece, Bulgaria, Rumania, Czechoslovakia, Poland, the 
Baltic States, Russia, and Scandinavia. Americans have been more 
ruthless in exterminating, or attempting to exterminate, any preda- 
tory animal conflicting, or presumed to conflict, with human interests. 
Unfortunately, the American method of dealing with predators by 
means of poison has attained a certain vogue in Bulgaria. 

A few of the more important vanishing mammals of Europe may 
be mentioned here. The Brown Bear (Ursus arctos) and the Wolf 
(Cam's lupus) are probably doomed to disappear almost entirely 
from Western Europe, although they will long survive in Russia 
and Siberia. The European Wildcat (Felis silvestris silvestris) has 
become extremely scarce in general; perhaps its greatest danger 
lies in extinction by dilution through interbreeding with feral 
Domestic Cats. The insular Wildcats (Cretan, Sardinian, Corsican, 
and British Felis agrius, F. sarda, F. reyi, and F. silvestris 
grampia) are probably endangered in like manner. The European 
Beaver (Castor fiber), persecuted for its fur, remains in only a few 
isolated colonies. There is some doubt as to whether any repre- 
sentatives of the Finland Reindeer (Rangifer tarandus fennicus) 
and the Novaya Zemlya Reindeer (R. t. pearsoni) still survive; 
the animal of Novaya Zemlya has fallen victim to visiting 
ships' crews and to Samoyed immigrants. While the stock of the 
Lithuanian Bison (Bison bonasus bonasus) is greatly reduced, and 
while there has been considerable mixture in captivity with the 
Caucasian Bison (B. b. caucasicus) and with the American Bison 
(Bison bison bison), energetic protection in sanctuaries assured 
its survival up to 1939, at least. Two of the four races of the 
Spanish Ibex (Capra pyrenaica) have been exterminated by exces- 
sive hunting, and the fate of those remaining has become uncertain 
during recent events in Spain. The Cyprian Mouflon (Ovis ophion 
ophion) has become reduced to a precariously small stock. 

The British Isles have long since lost the Brown Bear, the Wolf, 
the Beaver, the Wild Boar (Sus scrofa) , and the Reindeer (Rangifer 
tarandus) . No doubt insularity has here played a part in the early 
disappearance of these mammals. 



As long as the African Continent was occupied by primitive 
savages, without modern weapons, animal life was, in a large 
sense, in a virtual state of equilibrium. When European settle- 
ment began, and firearms were introduced, the death knell of a 
very considerable proportion of the population of large mammals 
was sounded. Thus the Atlas Bear (Ursus crowtheri) , the Barbary 
and the Cape Lions (Leo leo leo and L. I. melanochaitus) , the 
Quagga (Hippotigris quagga) , Burchell's Zebra (Hippotigris bur- 
chellii burchellii), the Bubal Hartebeest (Alcelaphus busclaphus 
buselaphus) , the Rufous Gazelle (Gazella rufina) , and the Blaauw- 
bok (Hippotragus leucophaeics) have departed finally and completely 
from the African scene. The typical subspecies of the Cape Harte- 
beest (Alcelaphus caama) may also be extinct, but imperfect knowl- 
edge of its distribution precludes a definite statement. A long 
time previously the Algerian Wild Ass (Asinus atlanticus) became 
extinct, from unknown causes. These losses by extinction are 
divided almost equally between South Africa the region most 
thoroughly settled by Europeans and the Barbary States, where 
the well-armed Moors long held sway. 

A century ago the Boer hide-hunters decimated the remarkable 
antelope and zebra fauna of South Africa. In the last half-century, 
firearms in the hands of improvident and short-sighted natives have 
wrought extremely serious havoc among the dwindling herds of 
African game in general. As intertribal warfare has practically 
ceased, and as the benefits of modern medicine and sanitation have 
penetrated far into the jungles and deserts, the native populations 
have increased, and their demands for a meat diet have decimated 
the game. Encircling fires, a method of hunting practiced on a 
fairly large scale in the savanna regions, have been extremely 
destructive, even in the absence of firearms. Professional hunters 
in the employ of great industrial enterprises, as in various parts of 
the Belgian Congo, have simply wiped out the antelopes over large 
areas. Hasty and probably ill-considered campaigns for the control 
of the tsetse fly have too often resulted in hecatombs of the large 
game mammals. In recent years the animals of the desert, such as 
Oryx and Gazelles, have become subject to attack from motor cars. 

In South Africa the Bontebok (Damaliscus dorcas) , the Blesbok 
(Damaliscus phillipsi) , and the White-tailed Gnu (Connochaetes 
gnou) no longer roam the free veldt, but have become restricted to 
enclosed farms and preserves. A remnant of the Cape Mountain 
Zebra (Hippotigris zebra zebra) was preserved at the eleventh hour. 


Among other vanishing or threatened African mammals, the fol- 
lowing may be mentioned in particular: 

Barbary Lynx (Caracal caracal algirus) 

South African Bush Elephant (Loxodonta africana ajricand) 

African Manatee (Trichechus senegalensis) 

Nubian Wild Ass (Asinus asinus africanus) 

Somali Wild Ass (Asinus asinus somaliensis) 

Southern White Rhinoceros (Ceratotherium simum simum) 

Northern White Rhinoceros (Ceratotherium simum cottoni) 

Pygmy Hippopotamus (Choeropsis liberiensis) 

Barbary Stag (Cervus elaphus barbarus) 

Congo Giraffe (Giraffa camelopardalis congoensis) 

Nigerian Giraffe (Giraffa camelopardalis peralta) 

Angola Giraffe (Giraffa camelopardalis angolensis) 

Southern Giraffe (Giraffa camelopardalis capensis) 

Okapi (Okapia johnstoni) 

Cape Buffalo (Syncerus caffer cafjer, here restricted to the South African 


Egyptian Arui (Ammotragus lervia ornata) 
Libyan Arui (Ammotragus lervia fassini) 
Nubian Ibex (Capra nubiana nubiana) 
Abyssinian Ibex (Capra walie) 
Cuvier's Gazelle (Gazella cuvien) 
Slender-horned Gazelle (Gazella leptoceros) 
Mhorr Gazelle (Gazella dama mhorr) 
White Oryx (Aegoryx algazel) 
Giant Sable Antelope (Hippotragus variant) 
Addax (Addax nasomaculatus) 
Nyala (Tragelaphus angasii) 
Mountain Nyala (Tragelaphus buxtoni) 

Senegambian Giant Eland (Taurotragus derbianus derbianus) 
Congo Giant Eland (Taurotragus derbianus congolanus) 

A very considerable number of game reserves have been estab- 
lished in various parts of Africa, and there should be a great many 
more of them, effectively supervised. Herein lies the chief hope for 
the survival of many of the larger African mammals. 


The mammalian fauna of this great island is particularly note- 
worthy for its very high degree of endemism and for the prepon- 
derance of lemurs. Madagascar and its outliers boast no less than 
three families and forty species and subspecies of lemurs, not one 
of which extends to the African mainland. Fortunately a fair pro- 
portion of these remain more or less common, being protected 
from persecution by native superstition. However, one species, the 
Hairy-eared Mouse Lemur (Cheirogaleus trichotis) , is apparently 
extinct. The following seem to exist in very small numbers, and 


should be safeguarded by every possible means from further de- 

Coquerel's Dwarf Lemur (Microcebus coquereli) 
Crossley's Mouse Lemur (C heirogaleus major crossleyi) 
Gray Lemur (Hapalemur griseus griseus) 
Broad-nosed Gentle Lemur (Hapalemur simus) 
Diademed Sifaka (Propithecus diadema diadema) 
Major's Sifaka (Propithecus verreauxi majori) 
Aye-aye (Daubentonia madagascariensm) 

Two peculiar carnivores, the Fossane (Fossa fossa) and the Fossa 
(Cryptoprocta ferox) , are also endemic in Madagascar. The former 
is accorded protection under Schedule A of the London Convention 
of 1933, and probably the latter is almost equally deserving of con- 

Perhaps the greatest danger to mammalian life in Madagascar is 
the steady reduction of the forest areas through burning and clear- 
ing by the natives. It is highly important from the point of view 
of conservation that this process should be halted. 


It may be of interest to the historian of mammalogy to list the 
extinct forms here in some sort of chronological order. They will be 
arranged chiefly by half-century periods and by regions within those 
periods; but those forms that passed out of existence prior to 1800 
will be placed in a single group. It should be borne in mind that 
in most cases the date of extinction can be only roughly indicated. 
For this reason the sequence within the regional half-century groups 
will be systematic rather than chronological. In some cases, how- 
ever, it is possible to add a more approximate date of extinction 
after the name of the species or subspecies. Certain cases of probable 
but unproved extinction are indicated by a question mark. 

Years 1-1800 (33 forms) : 


European Lion (Leo leo subsp.), 80-100 
European Wild Horse (Equus caballvs subsp.) 
Aurochs (Bos primigenius) , 1627 


Algerian Wild Ass (Asinus atlanticus) 
Blaauwbok (Hippotragus leucophaeus] , 1800 



Four Antillean insectivores (Nesophontes edithae; N. micrus; N. 

longirostris ; N. zamicrus) 
Lesser Falcate-winged Bat (Phyllops veins] 
Cuban Yellow Bat (Natalus primus) 

Smaller Puerto Rican Ground Sloth (Acratocnus odontrigonus) 
Larger Puerto Rican Ground Sloth (Acratocnus major) 
Smaller Hispaniolan Ground^loth ( Acratocnus (?) comes) 
Larger Hispaniolan Ground cloth (Parocnus serus) 
Barbuda Musk-rat (Megalomys audreyae) 

Hispaniolan Spiny Rat two species (Brotomys voratus; B. contractus) 
Cuban Short-tailed Hutia (Geocapromys columbianus) 
Crooked Island Hutia (Geocapromys ingrahami irrectus) 
Great Abaco Hutia (Geocapromys ingrahami abaconis) 
Haitian Hexolobodon (Hexolobodon phenax) 
Least Hispaniolan Hutia (Plagiodontia spelaeum) 
Puerto Rican Isolobodon (Isolobodon portoricensis) 
Haitian Isolobodon (Isolobodon levir) 
Narrow-toothed Hutia (Aphaetreus montanus) 
Two agoutilike rodents (Heteropsomys insularis; Homopsomys antil- 


A Puerto Rican hystricomorph (Heptaxodon bidens) 
"Quemi" of Oviedo (Quemisia gravis), about 1550? 
A Puerto Rican giant rodent (Elasmodontomys obliquus) 

Patagonian Giant Ground Sloth (Grypotherium listai) 

Steller's Sea-cow (Hydrodamalis gigas), 1768 

Years 1801-1850 (2 forms) : 

Eastern Bison (Bison bison pennsylvanicus) , 1825 


Hispaniolan Hutia (Plagiodontia aedium) 
Years 1851-1900 (31 forms) : 

Gilbert's Rat-kangaroo (Potorous gilbertii) 

i Possibly the extinction of some of the forms listed under this heading, known 
from bones found in cavern deposits, may have occurred more than 2,000 years 
ago. They are recognized, however, as pertaining to the Recent fauna. A. W. 


Portuguese Ibex (Capra pyrenaica lusitanica) , about 1892 


Atlas Bear ( Ursus crowtheri) 

Cape Lion (Leo leo melanochaitus) , about 1865 

Quagga (Hippotigris quagga), about 1878 

Hairy-eared Mouse Lemur (Cheirogaleus trichotis) 


Gull Island Meadow Mouse (Microtus pennsylvanicus nesophilus), 


Plains Grizzly (Ursus horribUis horribUis) 
California Coast Grizzly (Ursus calif ornicu*) , about 1886 
Sacramento Grizzly (Ursus colusus), about 1862 
Navajo Grizzly ( Ursus texensis navaho) 
Sonora Grizzly (Ursus kennerleyi) 
Mendocino Grizzly (Ursus mendocinensis) , about 1875 
New Mexico Grizzly (Ursus horriaeus) 
Sea Mink (Mustela macrodon), about 1880 
Eastern Wapiti (Cervus canadensis canadensis), about 1885 
Oregon Bison (Bison bison oregonus), about 1850's 


?Two Antillean insectivores (Nesophontes paramicrus; N. hypomicrus) 

?Puerto Rican Long-nosed Bat (Monophyllus frater) 

? Jamaican Long-tongued Bat (Reithronycteris aphylla) 

A Puerto Rican bat (Stenoderma rufum) 

?Puerto Rican Long-tongued Bat (PhyUonycteris major) 

?Haitian Long-tongued Bat (PhyUonycteris obtusa) 

Jamaican Rice Rat (Oryzomys antUlarum), about 1880's 

St. Vincent Rice Rat (Oryzomys victus), about 1897? 

Santa Lucia Musk-rat (Megalomys luciae) 

Larger Cuban Spiny Rat (Boromys offella) 

Lesser Cuban Spiny Rat (Boromys torrei) 

Antarctic Wolf (Dusicyon australis), 1876 

Chatham Island Rice Rat (Oryzomys galapagoensis) 


Years 1901-1944 (40 forms) : 


Freckled Marsupial Mouse (Antechinus apicalis) 

New South Wales Barred Bandicoot (Perameles jasciatd) 

Western Barred Bandicoot (Perameles myosura myosura) 

Nalpa Bilby (Macrotis lagotis grandis) 

Leadbeater's Opossum (Gymnobelideus leadbeateri) 

Gaimard's Rat-kangaroo (Bettongia gaimardi) 

Broad-faced Rat-kangaroo (Potorous platyops) 

Parma Wallaby (Thylogale parma) 

Toolach Wallaby (WaUabia greyi) 

White-tailed Rat (Zyzomys argurus ar gurus) 


Christmas Island Shrew (Crocidura juliginosa trichura), about 1904 
Maclear's Rat (Rattus macleari), about 1904 
Bulldog Rat (Rattus nativitatis) , about 1904 


Japanese Wolf (Canis hodophttax) 

Syrian Wild Ass (Asinus hemionus hemippus), about 1927 

Schomburgk's Deer (Rucervus schomburgki) , 1930's 


Caucasian Bison (Bison bonasus caucasicus), 1930's 
Pyrenean Ibex (Capra pyrenaica pyrenaica), 1910's 


Barbary Lion (Leo leo leo), 1922 

Burchell's Zebra (Hippotigris burchellii burchellii) 

Bubal Hartebeest (Alcelaphus buselaphus buselaphus), 1920's? 

Rufous Gazelle (Gazella rufina), 1920's? 


Long-eared Kit Fox (Vulpes macrotis macrotis), 1900's 

Newfoundland Wolf (Canis lupus beothucus), 1910's 

Florida Wolf (Canis niger niger), 1920's 

Tejon Grizzly (Ursus tularensis), 1916 

Texas Grizzly (Ursus texensis texensis), 1910's? 

?Mount Taylor Grizzly (Ursus perturbans) 

Black Hills Grizzly (Ursus rogersi bisonophagus) 

?Lillooet Grizzly (Ursus pervagor) 

?Klamath Grizzly (Ursus klamathensis) 

Southern California Grizzly (Ursus magister), 1908 


?Apache Grizzly (Ursus apache) 

Henshaw's Grizzly (Ursus henshawi), 1920's 

Eastern Cougar (Felis concolor couguar] 

Arizona Wapiti (Cervus canadensis merriami), 1906 

Badlands Bighorn (Ovis canadensis auduboni), 1900's? 


Cuban Solenodon (Solenodon cubanus), about 1910 
Martinique Musk-rat (Megalomys desmarestii), 1902 

James Island Rice Rat (Nesoryzomys swarthi) 

This record shows a steadily accelerated rate of extinction in 
each of the last three half-century periods. About 38 percent of 
the losses have been sustained since 1900. This indicates how diffi- 
cult is the task of preserving native faunas in the present era of 
intensive modern invention and industrial expansion. 


The following record indicates how these losses by extinction are 
divided among the various mammalian families: 

Bears (Ursidae), 17 

Spiny rats and their relatives (Echimyidae), 15 

Cattle, sheep, goats, and antelopes (Bovidae), 10 

Hamsterlike rodents (Cricetidae), 8 

Antillean insectivores (Nesophontidae), 6 

Leaf -nosed bats (Phyllostomidae), 6 

Kangaroos and their relatives (Macropodidae), 5 

Wolves and foxes (Canidae), 5 

Horses, zebras, and asses (Equidae), 5 

Ground sloths (Megalonychidae), 4 

Cats (Felidae), 4 

Bandicoots (Peramelidae), 3 

Old World rats (Muridae), 3 

Deer (Cervidae), 3 

Giant rats (Dinomyidae), 2 

Dasyures and their relatives (Dasyuridae), 1 

Phalangers and their relatives (Phalangeridae), 1 

Solenodons (Solenodontidae), 1 

Shrews (Soricidae), 1 

Long-legged bats (Natalidae), 1 

Lemurs (Lemuridae), 1 

Giant ground sloths (Megatheriidae), 1 

Heptaxodon (Heptaxodontidae), 1 

Weasels and their relatives (Mustelidae), 1 

Steller's Sea-cow (Hydrodamalidae), 1 

There is the clearest sort of significance in the losses sustained 
by the larger predatory mammals as a group (Ursidae, Canidae, 


and Felidae) , because of their competition with man for food in the 
shape of the ungulate mammals, both wild and domesticated, such 
as cattle, sheep, goats, antelopes, horses, asses, swine, and deer. 
In the case of such formidable carnivores as wolves, bears, lions, 
tigers, and leopards, the matter of outright self-defense on man's 
part may also be involved. Moreover, it is natural that the large 
game species of the cattle and deer families, which require extensive 
feeding grounds and are eagerly sought by mankind for food, 
should have suffered some of the principal losses. 


During the past 2,000 years the world has lost, through extinc- 
tion, about 106 forms (species or subspecies) of mammals. They 
are distributed by regions as follows: Australia, 11; Malay Archi- 
pelago, 3; Asia, 3; Europe, 6; Africa, 9; Madagascar, 1; North 
America, 27; West Indies, 41; South America, 1; Falkland Is- 
lands, 1; Galapagos Islands, 2; oceans, 1. Approximately 67 per- 
cent of these losses have occurred during the past century, and 38 
percent during the past half-century. Thus the rate of extinction 
is being steadily accelerated. 

In addition to the mammals already extinct, more than 600 others 
require consideration as vanishing or threatened forms. 

Insular faunas, partly by reason of their circumscribed nature 
and partly by reason of a certain lack of adaptability or self- 
defense, are particularly vulnerable to attack or competition by man 
and by certain mammalian pests introduced by him. There may be 
a further reason for the decadence of insular faunas in some cases, 
such as that of the West Indies, in the virtually total lack of native 
mammalian predators; these would doubtless have played a bene- 
ficial role by eliminating the less fit individuals, and thereby con- 
tributing to the survival of the fittest individuals, among the species 
preyed upon. 

In general, it is fairly obvious that species of restricted distribu- 
tion and specialized habits have less chance of survival than those 
of wide distribution and generalized habits. 

The primary factor in the depletion of the world's mammalian 
faunas is civilized man, operating either directly through excessive 
hunting and poisoning, or indirectly through invading or destroying 
natural habitats, placing firearms in the hands of primitive peoples, 
or subjecting the primitive faunas of Australia and of various 
islands to the introduction of aggressive foreign mammals, including 
fox, mongoose, cat, rat, mouse, and rabbit. Except in the West 
Indies, comparatively few species seem to have died out within the 
past 2,000 years from natural causes, such as evolutionary senility, 
disease, or climatic change. 


The chief hope for the survival of the larger mammals of the 
world lies in the establishment and maintenance of a sufficient num- 
ber of sanctuaries. This will avail in most parts of the world, but 
the matter is not so simple in Australia. Unless sanctuaries in that 
country can be surrounded with fences that are proof against foxes, 
rabbits, cats, and house rats, even they will not avail for many of 
the smaller Australian mammals. So perhaps the darkest picture 
today, as far as the future of mammals is concerned, is to be found in 
Australia, where many of the primitive native species cannot stand 
up against the highly organized introduced pests, and where condi- 
tions have gotten largely beyond human control. 


Order MARSUPIALIA: Marsupials 

Family DASYURIDAE: Dasyures, etc. 

This family, consisting of about 13 genera, is limited to Australia, 
Tasmania, New Guinea, and certain smaller neighboring islands. 
Of approximately 75 known forms, 14 call for discussion in the 
present work. 

Freckled Marsupial Mouse 


Phascogale apicalis J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 9, p. 518, 1842. 

("Doubtless from Australasia" =. South-Western Australia, fide Iredale and 

Troughton, 1934, p. 6.) 
FIGS.: Gould, 1863, vol. 1, pi. 39; Cabrera, 1919, pi. 5, fig. 2. 

This marsupial of Western Australia, little larger than a mouse, 
does not seem to have been collected for more than 30 years and is 
probably extinct. 

The general color is freckled reddish gray ; eye ring whitish ; under 
parts dull white or yellowish; pouch hairs dark rufous; front and 
outside of forearm rufous; rest of outer surface of limbs dull gray; 
ears short; tail tapering, variegated like back, and tipped with black. 
Head and body, 111-120 mm.; tail, 85-89 mm. (Thomas, 1888, 
pp. 277-278.) 

Very little information concerning this species is on record, and 
some of that is conflicting. Gould evidently gave it much too wide 
a range in stating (1863, p. 46) that it "is very generally distributed 
over every part of the colony of Western Australia." His actual 
records are from the vicinity of Moore's River, Perth, and King 
George's Sound. 

Thomas (1888, p. 278) records specimens from Albany and Vic- 
toria Plains, Western Australia, and even from Queensland. 

Shortridge states (1910, p. 840; map, p. 842) that it is "confined 
to the forest districts of the South-W T est, where it is apparently a 
rare species." He records three specimens from Albany in the Perth 



On the other hand, Glauert states (1933, p. 19) that the species 
is not represented in the West Australian Museum at Perth and is 
now probably extinct. 

Furthermore, despite the various specimens recorded by Gould, 
Krefft, Thomas, and Shortridge, E. Le G. Troughton writes (in litt., 
April 16, 1937) that "this species is not represented in the Perth 
Museum, and is probably known only from the type in the British 
Museum, and one in the Australian Museum. Apparently extinct or 
represented by small colonies only." Iredale and Troughton 
(1934, p. 6) limit the range to "South -Western Australia," omitting 

The species was probably either neutral or beneficial in its habits, 
for Gilbert (in Gould, 1863, p. 46) found the remains of insects in 
the stomachs he examined. 

No particular reason for its extinction seems to have been sug- 
gested, but the generally adverse conditions now facing the smaller 
marsupials of Australia are doubtless sufficient to account for it. 

Large Brush-tailed Phascogale; Brush-tailed Pouched Mouse 


Viverra tapoataja Meyer, Zool. Entdeck., p. 28, 1793. (Based upon "The 

Tapoa Tafa" of White, Jour. Voy. New South Wales, p. 281, pi. 58, 1790; 

type locality, Sydney, New South Wales.) 
SYNONYM: Didelphis penicillata Shaw (1800). 
FIGS.: Waterhouse, 1841, pi. 8; Gould, 1845, vol. 1, pi. 31; Lydekker, 1894, 

pi. 28; Jones, 1923, p. 99, fig. 60; Le Souef and Burrell, 1926, fig. 93; 

Fleay, 1934, pis. 19, 20. 

Though very considerably reduced in numbers, this animal still 
maintains itself in various localities through its wide range over 
the southern parts of Australia. 

Form stout and strong; general color finely grizzled pale gray; 
muzzle with indistinct darker stripe; ears very large, thin, nearly 
naked; under parts white or pale gray; pouch hairs dull rufous, 
tipped with white; terminal three-fifths of tail with a thick black 
brush. Head and body, 240 mm.; tail, 225 mm. (Thomas, 1888, 
pp. 295-296.) 

The general range is "southern Australia, from south Queensland 
to Western Australia" (Iredale and Troughton, 1934, p. 7) . 

Though once a familiar animal to settlers whose homes were in the more 
wooded districts, P. penicillata is unknown to the rising generation of country 
people. ... It seems astonishing that so small an animal could ever have 
been a real menace to the poultry run of the settler, and yet it is credited 
with being a determined slayer of chickens, and one which killed not merely 
to appease its appetite. Many of the older residents in South Australia have 
caught the animal red-handed, and as with the Native Cat, it seems a re- 


markable thing that so well equipped a carnivore should have been reduced 
to a condition bordering on extinction in so comparatively short a time. 
What its range within the State may have been is difficult to determine. 
It was not met with by the Horn Expedition, but an animal which answers 
very much to its description, but of which no specimen is available, apparently 
exists over a wide area in the Centre. By the South Australian Murray River 
natives it was well known under the name of "Pundi" but it has not been 
seen in their district for very many years. (Jones, 1923, p. 101.) 

Shortridge comments (1910, p. 839; map, p. 841) on its status 
in Western Australia as follows: 

"Although not plentiful this species seems to have a more general 
range in South-Western Australia than the smaller Phascogales. 

"Doubtfully recorded from as far inland as Kalgoorlie, where it 
would probably be only a straggler. . . . 

"Occasionally frequenting the neighbourhood of farms, where 
according to natives they come after mice." 

According to Glauert (1933, p. 19), it occurs in the southwest of 
Western Australia, from Fremantle to the south coast and inland 
to Merredin. Twenty specimens had been received at the Perth 
Museum in the preceding five years. 

For many years since the ravages of disease during the years 1898-1900, 
that agile and courageous little killer, "the brush-tailed rat" of the bushman, 
has been very scarce in the majority of its old haunts in Victoria and New 
South Wales. . . . 

The black "bottle-brush" tail and coat of 'possum-grey fur, combined with 
the amazingly agile movements of this lithe rat-sized marsupial, at once excite 
admiration. However, few people have enjoyed the spectacle of the nocturnal 
and arboreal creature making its lightning movements up and down the 
Eucalypt trunks "corkscrewing" round the boles to elude observation, or 
else bounding lightly, like a squirrel, from tree to tree. (Fleay, 1934, p. 89.) 

In Victoria, according to C. W. Brazenor (in litt., March 3, 1937) , 
the animal is "holding its own and common in timbered country." 

Le Souef and Burrell (1926, pp. 333-336) give the following 
account: "Some species, notably the brush -tailed and the lesser 
brush-tailed phascogales, are now rather scarce over the greater 
part of their range, having been greatly reduced by disease, which 
swept off large numbers of native animals in 1898-9-1900. Cats 
have also been very destructive. . . . 

"This species is more carnivorous than most members of the 
family. Moreover, it is very useful, in that it seems especially to 
catch rats and mice. There are instances of it following up plagues 
of these rodents and doing a good deal toward thinning them out." 

E. Le G. Trough ton (in litt., April 16, 1937) regards it as an 
active and resourceful species, whose survival is apparently assured, 
at least in the mountainous regions of its range. 

[A northern subspecies (P. t. pirata Thomas, 1904; type locality, 
"South Alligator River," Northern Australia) ranges across the 


northern part of the continent, from the Dawson Valley, Queens- 
land, to the Kimberley Division of Western Australia. It "appears 
to be very rare on the Dawson but still has a good hold on the 
wetter coastal country of the Fitzroy" (Finlayson, 1934, p. 226). 
It is reported as numerous in Arnhem Land, Northern Australia 
(Le Souef and Burrell, 1926, p. 336) . Its range lies largely outside 
that of the introduced fox, and its chances of survival are probably 
better than those of the southern subspecies.] 

Red-tailed Phascogale; Lesser Brush-tailed Pouched Mouse 


Phascogale calurus Gould, Proc. Zool. Soc. London 1844, p. 104, 1844. ("In 
the interior of Western Australia" = the Military Station on Williams 
River, fide Gould, 1863, vol. 1, p. 39.) 

FIGS.: Gould, 1863, vol. 1, pi. 32; Waterhouse, 1846, pi. 14, fig. 2. 

The Red-tailed Phascogale, of South, Central, and Western Aus- 
tralia, is so rare that few more than a dozen specimens seem to have 
been placed on record. It is evidently a vanishing species. 

General color ashy gray; under parts creamy white; ears large, 
nearly naked except at base, where there are some yellow hairs; 
basal half of tail rusty red above, black below; terminal half bushy, 
black (Gould, 1844, p. 105) . Head and body, 125 mm.; tail, 147 mm. 
(Thomas, 1888, p. 297.) 

Shortridge writes (1910, p. 839; map, p. 840) : "Very rare, seem- 
ing hitherto to have been recorded only four times from Western 
Australia; once from the Williams River, where it was originally 
obtained by Gilbert, and three times since from around Kojonup." 
Glauert (1933, p. 19) gives its range in Western Australia as "Lower 
South-West from Narrogin to Kojonup"; he adds that it "seems to 
be rather rare, six specimens only having reached the [Perth] 
Museum within the last five years." 

For Central Australia Spencer (1896, p. 30) records only a single 
specimen, taken at Alice Springs, and remarks that it "is evidently 
not a common form in the' central district." 

"The measurements given in the British Museum Catalogue of 
1888 are taken from an Adelaide specimen, but I have failed to 
trace any recent records of the animal in South Australia. . . . 
To-day it is impossible to define its former range in the State, or, 
unfortunately, even to attest to its present existence." (Jones, 
1923, p. 102.) 

Gould's statement of the range (1863, vol. 1, p. xxvii) as the 
"interior of New South Wales and the colony of Victoria" is ob- 
viously incomplete and supported by rather meager evidence. How- 


ever, Krefft states (1871 [p. 40]) that the animal occurs in New 
South Wales, near the Darling River, and Iredale and Troughton 
(1934, p. 8) include Victoria in the range. 

On Williams River, Gilbert (in Gould, 1863, vol. 1, p. 39) records 
the species as invading a storeroom. The type specimen was cap- 
tured in that locality by a Domestic Cat. 

"Some species, notably the brush-tailed and the lesser brush- 
tailed phascogales, are now rather scarce over the greater part of 
their range, having been greatly reduced by disease, which swept 
off large numbers of native animals in 1898-9-1900. Cats have also 
been very destructive." (Le Souef and Burrell, 1926, p. 333.) 

Here we seem to have yet one more melancholy case of the virtual 
disappearance of a species before any adequate knowledge of its life 
history or even of its distribution was obtained. 

Slender-tailed Pouched Mouse; Gray Pouched Mouse; 

"Common" Pouched Mouse; Slender 



This animal seems to have disappeared over considerable por- 
tions of its original wide range in Australia. It has been divided 
into the following four subspecies: 


Phascogale murina Waterhouse, Proc. Zool. Soc. London 1837, p. 76, 1838. 

("North of Hunter's River, New South Wales.") 
FIGS.: Waterhouse, 1841, pi. 10; Gould, 1863, vol. 1, pi. 43; Lydekker, 1894, 

pi. 29 (subsp.?) ; Le Souef and Burrell, 1926, fig. 94. 

Fur short and soft ; general color above gray, with a faint yellow- 
ish tint; feet, under parts, and face beneath eyes white; tail covered 
with minute silvery-white hairs. Head and body, 76 mm.; tail, 
65 mm. (Waterhouse, 1838, p. 76.) 

This form occurs in New South Wales and southern Queensland. 
Waterhouse reported it from north of Hunter's River, New South 
Wales, and Gilbert found it on the Severn River in the same state 
(Gould, 1863, vol. 1, p. 50). It was perhaps this form that Gould 
recorded (1863, vol. 1, p. 49 as Antechinus albipes) from the 
Darling Downs of New South Wales. 

This little marsupial, if sufficiently abundant, would evidently 
act as a check on one of the introduced rodent pests. "Mr. A. C. V. 
Bligh, of Toowoomba, Queensland, reports S. murina as being 
numerous at the same time as the common mouse (M. musculus) , 
and feeding upon the latter" (Le Souef and Burrell, 1926, p. 355) . 



Phascogale albipes Waterhouse, Proc. Zool. Soc. London 1842, p. 48, 1842. 

("Port Adelaide," South Australia.) 
Fia.: Gould, 1852, vol. 1, pi. 42. 

Upper parts brownish (the hairs being annulated with yellow 
near the tip and with black at the tip) ; hairs of under parts deep 
gray, tipped with white; feet white; tail dark, with very minute 
hairs. Head and body, 95 mm.; tail, 80 mm. (Waterhouse, 1842, 
p. 48.) 

This subspecies occurs in South Australia and Victoria. 

Gould gives it too wide a range in stating (1863, vol. 1, p. 49) 
that it "appears to be almost universally distributed over the whole 
of the southern coast of Australia, from Swan River to New South 

"These little animals . . . are caught in large numbers by the 
aborigines of the Murray" (Krefft, 1871, p. [41]). 

"Although in books this little animal passes uniformly under the 
name of 'Common' Pouched Mouse, it is by no means a common 
species. In South Australia it is not nearly so frequently met with 
as is S. crassicaudata. ... It is an animal which is very rarely 
seen except when it has fallen victim to a cat, and but little is 
known of its life history." (Jones, 1923, p. 118.) 

C. W. Brazenor (in litt., March 3, 1937) knows of no locality 
in Victoria in which this animal can be found at the present time. 


Antechinus juliginosus Gould, Mamm. Australia, vol. 1, pi. 41, 1852. ("At 
King George's Sound and in the vicinity of Perth," Western Australia. 
Thomas (1888, p. 305) lists the type specimen from "R. Avon, W. A.," 
which is in the general vicinity of Perth. Thus Iredale and Troughton 
(1934, p. 10) are evidently in error in giving, as the restricted type locality, 
"King George's Sound.") 

FIG.: Gould, 1852, pi. 41. 

Upper parts dark grayish brown, interspersed with longer black 
hairs; face lighter; a mark around the eyes black; chest sooty gray, 
with a narrow median line of buffy gray; rest of under parts pale 
grayish white; feet buffy white; tail dark reddish brown, grayish 
beneath. Head and body, 83 mm.; tail 83 mm. (Gould, 1863, vol. 1, 
p. 48.) 

The range is "South-Western Australia, more or less coastal, but 
inland to Katanning, Broomehill, Gnowangerup, and Bulong, near 
Kalgoorlie" (Glauert, 1933, p. 20) . 

Gould (1863, vol. 1, p. 48) considered it "very abundant, both 
at King George's Sound and in the vicinity of Perth." 


"These little animals ... are caught in large numbers by the 
aborigines ... of King George's Sound" (Krefft, 1871, p. [41]). 

Shortridge (1910, pp. 842-844; map, p. 843) speaks of it as 
"occurring throughout the South -West; appears to be more plenti- 
ful in the coastal districts wherever grass-trees (Xanthorrhoea) 
occur .... 

"On account of their habit of hiding among fallen timber or 
tree-stumps, the marsupial mice must invariably get exterminated 
wherever bush fires occur. This species, as well as Dromicia and 
the small Phascogales, has consequently become very scarce, espe- 
cially in the agricultural and more thickly populated areas. In addi- 
tion it is probably to a great extent killed off by the cats that have 
run wild in large numbers." 

More recently, however, Glauert (1933, p. 20) considers it still 
"a very common species in the South- West." 

Gilbert (in Gould, 1863, vol. 1, p. 48) found it insectivorous. 


Sminthopsis murina var. constricta Spencer, Kept. Horn Sci. Exped. Central 
Australia, pt. 2, zool., p. 33, 1896. ("Oodnadatta," South Australia.) 

General coloration similar to that of S. m. murina; foot broader; 
a small tuft of white hairs on posterior face of forearm; tail in- 
crassated. Head and body, 71 mm.; tail, 80 mm. (Spencer, 1896, 
p. 33.) 

In his original description of constricta, Spencer mentions only 
the single specimen from Oodnadatta, but on a previous page 
(1896, p. 32) he records a specimen of "S. murina" from Alice 
Springs, Central Australia, which perhaps belongs to the same 
form. He adds that the species "does not appear to be common 
in the central district." 

No additional information concerning the present subspecies 
seems to have come to light since its discovery was announced more 
than 45 years ago. 

Long-tailed Sminthopsis 


Sminthopsis longicaudatus Spencer, Proc. Royal Soc. Victoria, n. s., vol. 21, 
pt. 2, p. 449, 1909. ("West Australia.") 

This little animal is known from only a single specimen, from 
no more definite locality than "West Australia." 

General body color gray, tinged with rufous in parts; a darkish 
line through the eye; lips, chin, and feet white; tail scaly, with 


short stiff hairs. Head and body, 100 mm.; tail, 202 mm. (Spencer, 
1909, pp. 449-450.) 

"This species does not seem to have been recorded since its 
original discovery, and we have no information about the type 
locality beyond Spencer's vague 'West Australia' " (Glauert, 1933, 
p. 21). 

The apparently total lack of additional information concerning 
the species, during 30 years past, does not augur well for its present 
status, although there is always a possibility that it may have 
survived in some out-of-the-way corner of Western Australia. 

Common Eastern Native Cat; Viverrine Native Cat 


Didelphis Viverrina Shaw, Gen. Zool., vol. 1, pt. 2, p. 491, pi. Ill, 1800. (A 

composite species, based in part upon "The Tapoa Tafa" of White (Jour. 

Voy. New South Wales, p. 281, pi. 58, 1790) and in part upon "The 

Spotted Opossum" of Phillip (Voy. Botany Bay, p. 147, pi. 15, 1789). 

The name has become restricted to the latter; type locality, Botany Bay, 

New South Wales. Cf. Harper, 1940, p. 191.) 
FIGS.: Waterhouse, 1841, pi. 7 (as D. maugei) ; Gould, 1863, vol. 1, pi. 50; Krefft, 

1871, pi. 13; Lydekker, 1894, pi. 26; Le Souef and Burrell, 1926, figs. 87, 88; 

Fleay, 1932, pis. 3, 4; Pocock, 1937, p. 616, fig. 

This species, like its larger relative, Dasyurus maculatus, was 
distributed through eastern Australia and Tasmania and has suf- 
fered a similar or perhaps even greater reduction in numbers. 

There are two color phases, of which the black is the less com- 
mon. Fur thick and soft; general color either pale olive-gray or 
deep black, profusely spotted with white; belly and limbs paler 
than back; tail bushy, without spots, tipped with white in the gray 
phase. Head and body, 400-440 mm.; tail, 210-290 mm. (Thomas, 
1888, pp. 266-267.) 

The range includes New South Wales, Victoria, South Australia, 
and Tasmania. 

The history and status of the species are reviewed by Jones (1923, 
pp. 91-92) : 

It was abundant round, and even in the immediate precincts of, the larger 
Australian towns. Twenty years ago it was exceedingly common about Ade- 
laide. Still more recently it lived close to Melbourne; and to-day it is not 
uncommon in the suburbs of Sydney. . . . Very early in the days of colonisa- 
tion it was regarded with dislike because of the damage it did by killing 
poultry; but there are many settlers who would now welcome its return in 
order to keep the mice plagues within check. . . . 

There is no doubt that as a destroyer of mice, rats, and young rabbits the 
Native Cat played an extremely useful part in Australian rural economy, 
and despite the fact that it was an occasional robber of hen roosts its presence 
was a real asset to the country. 



Its range in South Australia was formerly very wide. On Kangaroo Island 
it appears to have been always more or less of a rarity. Thirty years ago it 
haunted the shores of the [Murray] river and lakes, being there very partial 
to a fish diet. To-day, if it exists at all in this State [South Australia], it 
must be an animal of the utmost rarity. Although there is no doubt that 
the influences which have been at work in the general process of the extermina- 
tion of the Australian fauna have operated to the full on the Native Cat; 
it is possible that another factor has come into play during the final scene 
of its passing. The animal has been trapped, poisoned and persecuted through- 
out the country .... The Native Cat, with its cunning and its activity, was 

FIG. 1. Common Eastern Native Cat (Dasyurus viverrinus) 

well able to look after itself, despite the fact that it was an extremely easy 
animal to trap. Its rapid decrease started about the year 1900, and during 
that and the two following years the so-called "common" Native Cat practically 
disappeared from South Australia. Much the same thing happened in Victoria 
and in New South Wales, with the exception of the district immediately 
round Sydney. It would seem that some epidemic disease must have spread 
through the Dasyures, and that after a lapse of twenty years the remnant 
has not succeeded in re-establishing itself. In the Animal Protection Act of 
1919 the Native Cat is not even mentioned. The evil or the good that it did 
has ceased to be a factor of any economic importance. 

At the present time it "still haunts the coastal cliffs and moun- 
tains about Sydney, also parts of Tasmania, Victoria, and New 
South Wales, where survival seems assured" (E. Le G. Troughton, 
in Hit., April 16, 1937) . 

In Victoria, according to C. W. Brazenor (in litt., March 3, 1937), 
it occurs in greatest numbers around Lake Corangamite but is also 


seen occasionally in the eastern part of the state. Further informa- 
tion is supplied by Fleay (1932, pp. 63-66) : 

From accounts supplied by people of these localities [about Lake Coranga- 
mite], the animals were not affected by the mysterious disease which an- 
nihilated many marsupials in other parts of the country in the first years of 
this century. Though still well known, however, they are becoming scarce, 
with the continuous work of rabbiters' dogs and traps, and the increase in 
settlement. . . . 

The adult males supported a host of parasites, and, when first brought to 
Melbourne, before being treated, they possessed numerous large ticks, sores 
infested with fly larvae, and the peculiar flea (Stephanocercus dasyuri) ; 
while investigations on Dasyures which had been caught in rabbit traps 
showed numerous nematode worms internally. 

On Kangaroo Island, South Australia, the species seems to have 
disappeared (Waite and Jones, 1927, p. 322) . 

In Tasmania it has fared better than in Australia. "The common 
Dasyure ... is to be met with in many localities in spite of the 
warfare waged against them in return for the toll they take of the 
settlers' poultry. In this respect, however, they cannot be con- 
sidered so destructive as the Tiger Cat' [D. maculatus]" (Lord 
and Scott, 1924, p. 270.) "The Dasyure is scattered throughout 
Tasmania, and still is very plentiful. This marsupial does not seem 
to either increase or decrease." (R. Boswell, in litt., May 13, 1937.) 

Despite Jones's statement (1923, p. 92) that "the animal's skin is 
of no commercial value," there evidently was a demand for it in 
former years. Lydekker writes (1894, p. 164) : "The fur being soft, 
the skins are suitable for linings; and from two to five thousand 
skins are annually imported into England. Formerly the grey skins 
fetched from about fivepence to sixpence each in the market, while 
the value of the black ones ranged from tenpence to a shilling. Of 
late years, however, there has been a fall in the price." 

The species is now under complete legal protection in Victoria 
and in Tasmania. 

Geoffrey's Native Cat; Black-tail CM! Native Cat 


Dasyurus Geoffroii Gould, Proc. Zool. Soc. London 1840, p. 151, 1841. ("Liver- 
pool Plains/' New South Wales.) 
FIGS.: Gould, 1851, vol. 1, pi. 51; Cabrera, 1919, pi. 4, fig. 2. 

The typical subspecies of Geoffrey's Native Cat seems to have 
become extinct over the greater part of its range, but it may sur- 
vive in parts of New South Wales and Queensland. 

General color fuscous, washed with yellow; head, back, and sides 
with white spots, smaller than those in D. viverrinus and D. macu- 


latus; tail long, terminal half black; under parts whitish. Head 
and body, 15 inches; tail, 11.5 inches. (Gould, 1841a, p. 151.) 

The former range of this animal extended from Victoria and 
South Australia through New South Wales to Queensland but 
apparently did not include the coast region of the southeast or the 
extreme north. 

According to Gould (1863, vol. 1, p. 58), the species (including 
both subspecies) "inhabits the whole of the southern portion of the 
country from Moreton Bay [Queensland] on the east to Swan 
River on the west." It "appears to be exclusively confined to the 
regions on the interior side of the hills, the specimens I have seen 
having been procured on the Liverpool Plains in New South Wales, 
the Murray Scrub in South Australia, and beyond the ranges of 
Swan River on the western coast." 

In Victoria it was always confined to the northwestern corner. 
The last known record was in 1857, and the species is now extinct 
in that state. (C. W. Brazenor, in Hit., March 3, 1937.) 

In South Australia there is no record other than that of Gould 
and a specimen listed in the British Museum Catalogue of 1888. 
"Men who have been professionally interested in the fauna of the 
State for a period of forty years are unaware of any examples being 
taken in South Australia proper. Unless it still lingers near to the 
northern limits of the State, it must probably be regarded as extinct 
in South Australia." (Jones, 1923, pp. 93-94.) 

In the Dawson Valley of Queensland, in 1905, it "was noticed to 
be suddenly numerous, but it completely vanished by 1906" (Fin- 
layson, 1934, p. 225) . It is represented in the Queensland Museum 
merely by two specimens without precise localities (Longman, 1930, 
p. 62). 

The eastern subspecies has very likely suffered in the same way 
as the western, which was "killed off as much as possible in the 
agricultural and more thickly populated districts on account of 
being so destructive to poultry" (Shortridge, 1910, pp. 838-839). 
Hoy (1923, p. 165) contributes information on an important enemy: 
"I . . . am told that domestic cats frequently kill and drag home 
adult native cats (Dasyurus viverrinus, D. geoffroyi, and D. hallu- 
catus)" Other possible causes underlying the sudden fluctuations 
in numbers of the species and its general disappearance over most 
of its range, have not been definitely explained. 

[The larger western subspecies (D. g. fortis Thomas) still occurs 
commonly in the southwest of Western Australia (Shortridge, 1910, 
pp. 837-839; Glauert, 1933, p. 18, and in litt., March 17, 1937). 
Some representative of the species perhaps fortis is reported from 
Central Australia but as nowhere common there (Finlayson, 19356, 


pp. 60-61). If this is the western subspecies, the eastern part of 
Western Australia forms a great blank in its known distribution.] 

Large Spotted-tailed Tiger-cat; Spotted-tailed Dasyure 


Viverra maculata Kerr, Anim. Kingdom of Linnaeus, p. 170, 1792. (Based 
upon the "Spotted Martin" of Phillip, Voy. Botany Bay, p. 276, pi. 46, 
1789; type locality "the neighborhood of Port Jackson" [Sydney], New 
South Wales.) 

FIGS.: Waterhouse, 1841, pi. 6 (as D. macrowrus} ; Gould, 1851, vol. 1, pi. 49; 
Lydekker, 1894, pi. 25; Raven, 1924, p. 25; Le Souef and Burrell, 1926, 
fig. 86; Fleay, 1932, p. 66, fig. 4, and pi. 5. 

This fierce and rather powerful animal, one of the largest of the 
carnivorous marsupials, is found in eastern Australia and in Tas- 
mania. Its range and its numbers have been reduced by settlement, 
though evidently not yet to the danger point. 

According to Phillip (1789, p. 276), the general color is black; 
body and tail irregularly blotched with white; tail tapering to a 
point; head and body, 18 inches; tail, nearly 18 inches. But Water- 
house (1846, pp. 440-441) and later authorities do not agree with 
Phillip and Kerr on the general color; it "varies from a very deep 
brown to a rich red-brown"; under parts "dirty yellow"; head and 
body, 17-24 inches; tail, 15-20 inches. 

The range includes "south-eastern Queensland, eastern New South 
Wales, Victoria, south-eastern South Australia, Tasmania" (Iredale 
and Troughton, 1934, p. 14) . Some of the earlier works extend the 
range into central or northern Queensland. According to Le Souef 
and Burrell (1926, p. 322), the species "is fairly common in Eastern 
Australia, from Cape York to Victoria." Half a century ago Thomas 
(1888, p. 265) considered it "approaching . . . complete extermina- 
tion in Australia"; but Ogilby (1892, p. 18) replied that it "is by 
no means uncommon nor seemingly has it any present intention 
of dying out in the mountainous and coastal districts of eastern 
Australia." On the Comboyne Plateau of New South Wales "it 
appears to be rather uncommon" (Chisholm, 1925, p. 72) . 

In Victoria it was "common in heavily scrubbed country till 
about 1907, at which time an epidemic of disease almost com- 
pletely destroyed the species. Has recovered somewhat in recent 
years and is found in some numbers in the Otway Ranges, and to a 
lesser extent scattered throughout the Dividing Range." (C. W. 
Brazenor, in litt., March 3, 1937.) "With the advent of settlement, 
disease, dogs, guns, traps, and . . . the fox, which exterminates 
the simple marsupial game of the Dasyure, we have come to the 
time, in Victoria, of the almost complete disappearance of these 


primitive carnivorous hunters" (Fleay, 1932, p. 68). The species 
now has complete legal protection in Victoria. 

"Probably never abundant in South Australia, the stronghold of 
the species was in the south-eastern portion of the State. It is 
possible that some few still exist in the less closely settled areas 
of the South-East." (Jones, 1923, p. 88.) 

In Tasmania it "is regarded as one of the settlers' greatest pests, 
owing to the toll it will take of his poultry" (Lord and Scott, 1924, 
p. 269). "The enemy of the settler's chickens, it is only natural 
that this species should be reduced in numbers, especially in the 
settled districts. Even so, this hardly accounts for the scarcity of 
this species in the more Southern Tasmanian localities in the last 
few years. In the North-West the species is still fairly common." 
(Lord, 1928, p. 22.) 

There are additional records of nocturnal raids on poultry on 
the mainland of Australia, and this habit naturally reacts against 
the species. "All three dasyures are doomed to extinction, since 
they are killed whenever met with by the man on the land" (Jones 
and Manson, 1935, p. 34) . "It is now being replaced by the domestic 
cat and the fox" (Raven, 1924, p. 25) . However, "it is able to kill 
wallabies and fairly large birds," and "one succeeded, after a 
severe battle, in killing a large tom-cat" (Le Souef and Burrell, 
1926, pp. 322-323). Although Lydekker wrote in 1894 (p. 160) that 
"its skin is but little valued by furriers," it must be remembered 
that many furs, formerly in little demand, now bring good prices. 

While the Dingo is generally considered responsible for the ex- 
tinction of the terrestrial Tasmanian Devil and Tasmanian Wolf in 
Australia, opinion seems divided as to whether it has seriously 
affected the status of the arboreal Spotted-tailed Tiger-cat on the 
mainland. E. Le G. Troughton (in litt., April 16, 1937) believes that 
this species, by reason of its furtive and aggressive disposition, 
should survive indefinitely in the dividing ranges of the east coast. 

Slender Native Cat; Slender Spotted-tailed Tiger-cat 


Dasyurus gracilis Ramsay, Proc. Linn. Soc. New South Wales, ser. 2, vol. 3, 
p. 1296, 1888. ("Bellenden-Ker Ranges," northern Queensland.) 

The Slender Native Cat is one of those species which, as far as 
known, has always been very rare. Apparently less than half a 
dozen specimens are on record all from northern Queensland. 

General color, above and below, deep blackish brown with white 
spots; tail spotted and closely furred, with a terminal tuft on the 
upper side. Total length, about 23 inches; tail, 9.3 inches. (Ramsay, 
op. tit., p. 1296.) 


The type specimen was collected by Robert Grant on the Bel- 
lenden-Ker Ranges, apparently in 1887. The species was next 
found years later by C. M. Hoy on the Atherton Tableland (Le 
Souef and Burrell, 1926, p. 324) . The late Henry C. Raven, of the 
American Museum of Natural History, informed me that he secured 
two or three specimens about 1922 in the same general region. 

Whatever the factors may be that seem to restrict so decidedly 
the numbers of the Slender Native Cat, they have not been ascer- 

According to E. Le G. Troughton (in litt., April 16, 1937) , the 
species is rarely captured because of the density of its mountain 
rain-forest habitat, and should therefore survive in parts of coastal 
Queensland for all time. 

Tasmanian Devil 


Didelphis ursina Harris, Trans. Linn. Soc. London, vol. 9, p. 176, pi. 19, 

fig. 2, 1808. ("Van Diemen's Land.") (Not Didelphis ursina Shaw (1800).) 
Ursinus harrisii Boitard, Jardin des Plantes, p. 290, "1842" = 1841. (Tasmania.) 
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mammif., vol. 7, pi. 113, 1842; 

Gould, 1851, vol. 1, pi. 48; Royal Nat. Hist., vol. 3, p. 271, fig., 1894-95; 

G. Smith, 1909, fig. 24; Raven, 1924, p. 25, fig., and 1929, p. 204, fig.; 

Le Souef and Burrell, 1926, fig. 85; Fleay, 1935, pi. 9; Pocock, 1937, 

p. 615, fig.; Reed and Lucas, 1937, p. 89, fig. 33. 

This fierce little beast occurred in past ages on the Australian 
mainland, where it presumably succumbed to the advancing Dingo. 
In 1912 a specimen, probably an escaped captive, was taken in 
Victoria. It "is now confined to Tasmania, where it maintains a 
rather precarious foothold in the wilder parts of the country" 
(Jones, 1923, p. 85). 

Whole body and upper part of tail covered with long coarse 
black hair; irregular blotches of white on shoulders, throat, or 
rump (G. P. Harris, 1808, p. 176) . It is a thickset, powerful animal, 
and, except for its tail, resembles a miniature bear in outline. Head 
and body, 670-825 mm.; tail, 258-300 mm. (Lord and Scott, 1924, 
p. 267.) 

"These animals were very common on our first settling at Hobart 
Town, and were particularly destructive to poultry, &c. They, 
however, furnished the convicts with a fresh meal, and the taste 
was said to be not unlike veal. As the settlement increased, and the 
ground became cleared, they were driven from their haunts near 
the town to the deeper recesses of the forests yet unexplored." (G. P. 
Harris, 1808, p. 177.) 

"The devil is destructive to sheep all over the colony, and is 



indeed the most destructive of our indigenous quadrupeds, the 
Thylacinus being much scarcer" (Gunn, 1838, p. 104). 

"It has now become so scarce in all the cultivated districts, that 
it is rarely, if ever, seen there in a state of nature; there are yet, 
however, large districts in Van Diemen's Land untrodden by man; 
and such localities, particularly the rocky gullies and vast forests 
on the western side of the island, afford it a secure retreat. . . . 

"In its disposition it is untameable and savage in the extreme, 
and is not only destructive to the smaller kangaroos and other 
native quadrupeds, but assails the sheep-folds and hen-roosts when- 
ever an opportunity occurs." (Gould, 1863, vol. 1, p. 55.) 

FIG. 2. Tasmanian Devil (Sarcophilus harrisii) 

"The Devil is far commoner than the Tiger and more widely dis- 
tributed through the island .... Like the Tiger it destroys sheep, 
making a single meal off each capture." (G. Smith, 1909, p. 97.) 

Lord (1928, p. 22) says of it: 

The Tasmanian Devil will probably survive for many years. Its hardy 
nature both in captivity and in its wild state cause [s] one to wonder how it 
came about that this species became extinct on the mainland within com- 
paratively recent geological times. It cannot be considered a pleasant animal 
to have much to do with, and numbers are killed by trappers in the course 
of their work. In the rougher sections of the country this species exists in 
fair numbers and there is every prospect of it remaining an inhabitant of 
such places for years to come. 

One or more Tasmanian Devils will often follow a Thylacine on its hunting 
excursions. The Thylacine will kill a wallaby or other small animal, select a 
few choice morsels, and pass on. The Devils will carry on the feast and 
consume the remnants, bones and all. 

According to R. Boswell (in Hit., May 13, 1937), it still exists 
in large enough numbers to enable it to be out of immediate danger 


of extinction. Civilization has been the great cause of its decreasing 
numbers. It has no legal protection. 

"In spite of his ungainly, ugly appearance, his whining snarls 
and unpleasant smell, the Tasmanian Devil is a creature of many 
amusing antics and distinctly unusual ways. Moreover, his position 
as the second largest of living marsupial carnivores, soon, perhaps, 
to be the largest when the rare Thylacine finally disappears, invests 
him with a peculiar interest." (Fleay, 1935, p. 100.) 

Tasmanian Wolf; Marsupial Wolf; Tasmanian Tiger; 



Didelphis cynocephala Harris, Trans. Linn. Soc. London, vol. 9, p. 174, pi. 19, 
fig. 1, 1808. ("Van Diemen's Land" [= Tasmania].) 

FIGS.: Waterhouse, 1841, pi. 5; Gould, 18*51, vol. 1, pis. 53, 54; Wolf, 1861, 
pi. 31; Krefft, 1871, pi. 12; Royal Nat. Hist., vol. 3, p. 270, fig., 1894-95; 
G. Smith, 1909, fig. 23; Cabrera, 1919, pi. 6; Australian Mus. Mag., vol. 1, 
no. 3, p. 62, frontisp., 1921; Le Souef and Burrell, 1926, fig. 84; Raven, 
1929, p. 207, fig.; Pocock, 1937, p. 614, fig.; Reed and Lucas, 1937, p. 85, 
fig. 31; Sharland, 1939, p. 23, fig. 

This largest and most formidable of living carnivorous marsupials 
is so seriously reduced in numbers that its fate seems to be hanging 
by a somewhat slender thread. 

General build doglike, but hind end tapering gradually to the 
tail; upper parts tawny grayish brown, with 16-19 blackish brown 
bands across the back, chiefly developed on the hind quarters; 
under parts paler. Head and body, 1230-1300 mm.; tail, 525-650 
mm.; height at shoulders, about 560 mm. (Chiefly from Lord and 
Scott, 1924, p. 264.) 

While a fossil form of Thylacine has been recorded from the 
Australian mainland, the range of the living form is restricted to 
Tasmania. The mainland Thylacine is presumed to have suc- 
cumbed as a consequence of the advent of the Dingo during the 
Pleistocene, for it probably could not compete successfully with 
that more highly organized animal. 

The Thylacine "is common in the more remote parts of the 
colony, and they are accordingly often caught at Woolworth and 
the Hampshire hills. . . . They are usually nocturnal in their 
attacks on sheep." (Gunn, 1838, p. 101.) 

It was with prophetic vision that Gould wrote long ago (1863, 
vol. 1, pp. 60-61) : 

When the comparatively small island of Tasmania becomes more densely 
populated, and its primitive forests are intersected with roads from the eastern 
to the western coast, the numbers of this singular animal will speedily diminish, 


extermination will have its full sway, and it will then, like the Wolf in 
England and Scotland, be recorded as an animal of the past: although this 
will be a source of much regret, neither the shepherd nor the farmer can be 
blamed for wishing to rid the island of so troublesome a creature. A price 
is already put upon the head of the native Tiger, as it is called; but the 
fastnesses of the Tasmanian rocky gullies, clothed with impenetrable forests, 
will, for the present, preserve it from destruction. 

. . . Although too feeble to make a successful attack on man, it commits 
sad havoc among the smaller quadrupeds of the country, and among the 
poultry, and other domestic animals of the settler; even sheep are not secure 
from its attacks .... 

"The damage which it inflicts on the flocks of the settlers has 
. . . given rise to a relentless war of extermination, which has 
resulted in the almost complete extinction of this, the largest of 
the Australasian Carnivores, in the more settled portions of the 
country" (Lydekker, 1894, p. 152). 

G. Smith (1909, pp. 96-97) wrote: 

The destructiveness of these animals is greatly enhanced by the fact that a 
Tiger will make only one meal of a sheep, merely sucking the blood from 
the jugular vein or perhaps devouring the fat round the kidneys, but it 
never returns to the same carcass. . . . The shepherds wage incessant war 
on the creature, in the summer laying traps and hunting it with dogs, in the 
winter following up its tracks through the snow. A reward of a ' pound is 
given for the head by the Government, but the shepherd generally rides 
round with the head to several sheep-owners in the district, and takes toll 
from them all before depositing it at the police station. In consequence a 
large reward must be offered for the carcass of a Tiger, and an offer of 10 
during a year for a live Tiger to be delivered in Launceston was unsuccessful. 
It pays the shepherd very much better just to hack off the head and take it 
round on his rides. Although the Tiger is by no means confined to the Lake 
District, it is more abundant here than anywhere else, though a stray individual 
may turn up on nearly all the big sheep stations throughout the island. 

Lord (1928, pp. 20-21) says of the Thylacine: 

The animal is confined practically to the rugged western portion of the 
island. From the more settled districts it has long since disappeared, and 
even in the more distant sheep runs it has been trapped out .... It is 
now also being killed out even in the rugged and more inaccessible parts 
of the country, which tends to reduce still further the remnants of this 
species. The explanation of this is that the Thylacine interferes with the 
trappers' snares. As a result, a powerful "springer" snare is set often in the 
vicinity of their "skinning yards," which are situated every quarter of a 
mile or so along the lines of snares. Thylacines or other animals caught 
in these powerful snares are, as a rule, too severely injured to be kept alive as 
specimens for zoological gardens, even if the trappers would take the trouble 
to bring them in. The extended trapping of recent years will tend, therefore, 
to restrict the Thylacine to the most rugged and unsettled portions of the 
West of the island. Here it may survive as a living species for years to come, 
but its eventual doom seems apparent unless such attempts as are being made 
at present by Mr. A. R. Reid (Curator of the Beaumaris Zoo, Hobart) to 
breed these animals in captivity are successful. . . . 

It is doubtful if the shy animal will breed within the confines of a Zoo, 
and it would be in the interests of science if a reserve could be set aside and 



netted in in order to prevent total extermination. ... If funds were available 
an area in the National Park might well be considered for such a reserve. 

"The Tasmanian Tiger is now only to be met with in a very few 
numbers. This animal is causing great concern in Tasmania at 
the present time. It is thought by many to be extinct, but this is 
not so. I have obtained authentic reports regarding its presence 
as having been seen as recently as January 1937 on the West 
Coast of Tasmania. 


FIG. 3. Tasmanian Wolf (Thylacinus cynocephalus) 

11 The former range of the Tasmanian Tiger must have been very 
great as I know of one Tasmanian, who with his brother, killed as 
many as twenty-four of these animals during one day, and received 
a reward of 1. 0. per head for each animal. 

"The Tasmanian Tiger is now wholly Protected." (R. Boswell, 
in litt., May 13, 1937.) 

"The significance of the mainland elimination, prior to settlement, 
of the largest living marsupial carnivore (Thylacinus) has already 
been noted, and latest reports from Tasmanian authorities indicate 
grave doubts for the insular survival of this unique example of 
parallelism" (Troughton, 1938, p. 408) . 

However, the latest news from Tasmania is distinctly encouraging. 
It comes in the form of a paper by Sharland (1939), which gives 
an account of several recent expeditions that have been sent to 
mountainous areas in the western part of the state by the Tasmanian 
Animals and Birds' Protection Board. From this account the fol- 
lowing information is derived: 

"The Thylacine exists to-day as but a remnant of the numbers 


which, 50 or 60 years ago, roamed the countryside, feeding on small 
marsupials and sheep. . . . Nowadays, certainly, it is rarely seen. 
. . . When the game season is opened every few years the animal is 
often caught in snares. But it is in no part specially common, and 
there are extensive areas in this region where it does not occur at 
all, or but sparsely, its distribution depending almost wholly on the 
presence of smaller 'game.' " (P. 20.) 

"The Thylacine has been known to attack dogs when cornered, 
but so far as I can determine there is no record of its ever having 
attacked man" (p. 32) . 

In a great amphitheatre about 25 to 40 miles in diameter, bounded 
by the King William, Prince of Wales, Norway, and other ranges, 
"we came upon many tracks made by Thylacine, indicating that the 
animal was fairly common and well distributed" (p. 34) . 

"The area enclosed by the mountains would make a splendid game 
sanctuary .... The Thylacine is probably as common here as in 
any other part of the West Coast." (P. 34.) 

Additional tracks were found in the Jane River region, where 
"the animal had apparently been trailing Wallaby" (p. 34). 

"No longer a menace to sheep-owners since its isolation in the 
remote parts of the State, the animal possesses a unique scientific 
value which is appreciated by the Board. While, up to half a 
century ago, it was fairly plentiful in the grazing country of the 
central plateau, and was known also to inhabit parts of the eastern 
tiers and other mountain forest areas adjacent to settlement, it has 
now practically disappeared from these districts, to make its last 
stand in the western section of the State." (P. 36.) The recent 
opening of a road through this remote region has had an adverse 
effect upon the Thylacine's prospects for survival. 

A mountainous area situated about Frenchman's Gap, east of 
Macquarie Harbor, and comprising approximately 300,000 acres, 
is suggested as a suitable sanctuary for the Thylacine and other 
animals (p. 38) . 

Family MYRMECOBIIDAE : Marsupial Anteaters 

The single genus of this family consists of two forms, both of 
which are treated here. They occur in the southern half of Australia. 

Banded Anteater; West Australian Numbat 


Myrmecobius fasciatus Waterhouse, Proc. Zool. Soc. London 1836, p. 69, 
1836. ("In the interior of the Swan River Settlement, about 90 miles to 
the S.E. of the mouth of that river," Western Australia.) 


FIGS.: Waterhouse, 1838a, pi. 27, and 1841, pi. 11; Gould, 1845, vol. 1, pi. 4; 
Lydekker, 1894, pi. 30; Royal Nat. Hist., vol. 3, p. 275, lower fig., 1894-95; 
Cabrera, 1919, pi. 7; Le Souef and Burrell, 1926, fig. 96. 

This beautiful little animal, representing a special family related 
to the dasyures, has been undergoing shrinkage of range and reduc- 
tion in numbers in Western Australia for a considerable period and 
is perhaps approaching extinction. 

Color above reddish ochre, interspersed with white hairs ; posterior 
half of body with alternate black and white bands; tail long-haired, 
mixed with black, white, and reddish ochre; legs chiefly pale buff; 
under parts yellowish white (Waterhouse, 1836, pp. 69-70). Form 
graceful, squirrellike ; a black stripe through the eye, and a white 
stripe above it. Head and body, 220-240 mm.; tail, 160-175 mm. 
(Thomas, 1888, pp. 311-314.) 

In earlier days its range extended west to the Darling Range, 
northwest to the vicinity of Moore's River, northeast to Laverton, 
east to Kalgoorlie and possibly to South Australia near the coast, 
and south to the vicinity of Albany (Shortridge, 1910, p. 846, map; 
Glauert, 1933, p. 22). Forty years ago it was "fairly numerous 
throughout the South-west, especially where the prevailing timbers 
are the white gum (Eucalyptus redunca) and the jam (Acacia 
acuminata) , getting less plentiful outside that area" (Shortridge, 
in Thomas, 1907, p. 772) . "The Western Australian animal is now 
excessively rare, and it is probable that before many years are 
passed it will follow its South Australian neighbour [M. f. rujus] 
into extinction" (Jones, 1923, p. 126). Troughton writes (1923, 
p. 155) that "this animal . . . can now only be found in a greatly 
restricted area"; he secured three specimens about 1921. It seems 
to survive chiefly in the southwestern corner of the state, between 
Perth and Albany. 

"The beautiful little Banded ant-eater is much sought after on 
account of its skin" (W. H. D. Le Souef, 1907, p. 406) . 

Le Souef and Burrell (1926, pp. 365-366) write: 

Quiet, inoffensive, without means of defence or offence, it is remarkable 
that the marsupial ant-eater has survived through the ages. This could 
happen only in Australia, where it did not come into competition with the 
more advanced forms of life. . . . 

It is abroad both by day and by night, and, being conspicuous and not at 
all speedy, it makes a fairly easy mark for predacious animals and birds, 
more especially the introduced cat and fox; to say nothing of the settlers' 
dogs. Consequently, it is one of the first animals to disappear before the 
inevitable opening up of the country, and it is now scarce over the greater 
part of its range. 

"The typical race ... is endangered by clearing, fires, and intro- 
duced pests, the advent of the fox alone probably spelling the ulti- 



mate doom of the terrestrial and non-burrowing highly specialized 
creature. . . . Myrmecobius may be regarded as one of the marsu- 
pials within sight of extermination, in this instance not due to 
exploitation by man, but as the result of settlement and introduced 
enemies. Hope for ultimate survival may rest with the introduction 
of a healthy colony to an island providing adequate supplies of 
favoured diet, and absence of enemies." (E. Le G. Troughton, 
in Hit., April 16, 1937.) 

Its "very existence ... is threatened by both fox and cat" 
(Troughton, 1938, p. 404) . 

South Australian Numbat; Rusty Numbat 


Myrmecobius rujus Jones, Mammals S. Australia, pt. 1, p. 123, figs. 79, 83, and 

84, 1923. ("South Australia.") 
FIG. : Jones, 1923, fig. 79. 

This form of Numbat, apparently extinct in New South Wales 
and coastal South Australia, still lingers in northwestern South 
Australia and in southwestern Central Australia. 

It differs from the West Australian Numbat in having the darker 
part of the lower back "a fine bright brown" instead of blackish; tail 
"a uniform grizzle of rust red and dark brown." Head and body, 
175 mm.; tail, 135 mm. (Jones, 1923, pp. 124-126.) Finlayson 
states (1933c, p. 204) that the outer surface of the ear is bright 
rufous instead of yellow and black, and he gives the following 
measurements for specimens from northwestern South Australia: 
head and body, 200-270 mm.; tail, 130-170 mm. 

"The New South Wales animal, reported fairly plentiful about 
the plains of the Murray and Darling Rivers in 1862, ... is 
apparently extinct" (E. Le G. Troughton, in litt., April 16, 1937). 

It is perhaps the present form to which Helms refers (1896, 
p. 255) in reporting the observations of the Elder Expedition 
somewhere in South or Western Australia: "A more exciting piece 
of work [by the natives] than digging for lizards is the excavating 
for the quick, little, banded anteater, Myrmecobius jasciatus, which 
animal often makes its lair over three feet below the surface." 
This expedition brought back a dried skin from the Everard Range, 
South Australia (Stirling and Zietz, 1893, p. 154) . 

Jones (1923, pp. 126-127) says of this Numbat: 

The Numbat was probably never a very abundant animal, but its distribu- 
tion was comparatively wide. Only twenty years ago it was met with along 
the scrub lands of the Murray, and earlier than that it existed quite near to 
Adelaide. Enquiries as to its present existence have produced negative replies 
from all those parts of the State in which there are schools, and the circulation 


of its picture and description to more remote districts have proved equally 
unavailing. The aboriginals who are attracted to civilisation, as it is represented 
by the East-West railway, know the animal, but so far have failed to supply 
any evidence as to its actual existence at the present time. If the Numbat 
still exists in South Australia it is probably towards the Western Australian 
border, and here it is probably the Western Australian form. The characteristic 
South Australian type has probably gone for ever. . . . The extermination 
of the Numbat is a tragedy in which man has probably played very little 
conscious part; it is no tale of ruthless slaughter for gain, such as is being 
rehearsed to-day in regard to the Australian fur-bearing animals, nor is it a 
case of determined persecution as is the case with the Tasmanian Devil. 
MyrmecobiiLS is an animal which is probably phylogenetically senile, which 

FIG. 4. South Australian Numbat (Myrmecobius jasciatus rujus). 
After Jones, 1923. 

has become highly specialised in function and degenerate in some details 
of structure. Added to this is the fact that its home is invariably made in 
the hollow of a fallen tree or a rotting log. Accidental bush fires and the 
intentional burning off of country seem to have found the Numbat an easy 
victim, and they have exterminated it as they are exterminating other small 
terrestrial Marsupials. There is no escape from a bush fire for the Numbat. 
It does not excavate deep burrows, it does not climb, it is not fleet of foot 
as its log home burns, it perishes. . . . 

It is surely a tragedy that this most interesting animal has probably passed 
out of existence in our State, and is rapidly repeating the process in a 
neighbouring one without any representative collection having been made of 
its remains. It will not be long before Myrmecobius will be as extinct as those 
Mesozoic Marsupials of the English Jurassic beds of which it has been said 
to be "actually an unmodified survivor." 

Since the publication of Jones's account, investigation by Finlay- 
son has shown that the species still survives in the arid center of 
the continent. Rewrites (1933c, p. 203) : 

"Its presence in the centre [in the Everard Range] was first 
established by the work of the Elder Expedition .... 


"Recent field work ... in the far north-west of this State [South 
Australia] (in a typical eremian environment) has shown . . . 
that Myrmecobius still has a wide distribution in the south-west 
parts of the centre beyond the limits of pastoral settlement, and in 
some localities is by no means uncommon. It is possible that these 
colonies actually link up with the far south-western ones in Western 
Australia in a continuous band of distribution." 

Finlayson here proposes (1933c, p. 203) to separate the central 
animal from that of Western Australia under the name of Myrme- 
cobius fasciatus var. rufus and gives (p. 204) as type locality 
"mulga sand dunes, south and south-west of the Everard Range, far 
north-west of State of South Australia." This name, however, is 
antedated by Myrmecobius rufus Jones (1923), which was intro- 
duced without any formal designation of type locality, but which 
was based upon "South Australian specimens, from the Murray 
and from near Adelaide" (Finlayson, 1933c, p. 205). The range, 
according to Finlayson (p. 204), is "at present apparently not 
north of about 25 S. lat., nor east of 132 30' E. long. To the south 
and west as yet undetermined. Formerly as far south as Adelaide, 
and probably ranging east into the Victorian and New South Wales 
mallee areas." 

Family PERAMELIDAE : Bandicoots 

The range of the bandicoots extends over Australia, Tasmania, 
New Guinea, and certain adjacent islands. There are about 9 
genera, represented by about 44 forms. Of the latter, accounts of 
12 appear in the following pages. 

Eastern Barred Bandicoot ; New South Wales Barred Bandicoot 


Perameles fasciata J. E. Gray, in Grey, Two Expeditions Australia, vol. 2, 
appendix, pp. 401, 407, 1841. ("Liverpool Plains and South Australia"; 
type locality restricted by Thomas (1922, p. 144) to "Liverpool Plains," 
New South Wales.) 

Fia: Gould, 1849, vol. 1, pi. 8. 

This bandicoot occurred formerly in New South Wales and 
Victoria. It has not been recorded for many years, however, and 
is probably extinct (A. S. Le Souef, in litt., February 15, 1937). 

This species has been more or less confused in descriptions with 
P. myosura notina. "Grey brown, rump with three black bands; 
tail white, with a black streak along the upper side. . . . Smaller 
than P. Gunnii." (J. E. Gray, in Grey, 1841, p. 407.) Upper parts 
penciled with black and yellow; sides yellow; under parts and feet 



white (Gould, 1863, vol. 1, p. 12). Outer surface of ears flesh color 
basally, darker terminally; sides of rump with four pale vertical 
bands running downward from near the middle line, the spaces 
between them brown or black (Thomas, 1888, p. 248) . 

"This elegant species . . . enjoys a wide range over the eastern 
. . . portions of Australia, but is more frequently met with in the 
country within the ranges . . . than in the districts lying between 
the mountains and the sea. In New South Wales, the stony ridges 
which branch off from the ranges towards the rivers Darling and 
Namoi, are localities in which it may always be found." (Gould, 
1863, vol. 1, p. 12.) 

IE. . 

FIG. 5. Eastern Barred Bandicoot (Perameles jasciata). After Gould, 1849. 

The animal is "now believed extinct though once well distributed 
over western Victoria and N. S. Wales. The ultimate fate of these 
small non-burrowing forms is most uncertain." (E. Le G. Trough- 
ton, in litt., April 16, 1937.) 

Tasmania ii Barred Bandicoot; Gunn's Striped Bandicoot 


Perameles Gunnii J. E. Gray, Ann. Nat. Hist., ser. 1, vol. 1, p. 107, 1838. 

("Van Diemen's Land" [= Tasmania].) 
FIGS.: Waterhouse, 1841, pi. 15; Gould, 1859, vol. 1, pi. 9; Lydekker, 1894, 

pi. 21. 

While this species still occurs in numbers in Tasmania, it is 
"bordering on extinction in Victoria" (David H. Fleay, in litt., 
June 1,1937). 

Muzzle tapering, gray-brown; under parts, feet, tail, and four 


broad bands on each side of the rump white (J. E. Gray, in Gunn, 
1838, pp. 107-108). General color grizzled yellowish brown; outer 
surface of ears yellowish brown, with a darker terminal blotch; 
sides of rump with four more or less distinct pale vertical bands 
running downward from near the middle line, the spaces between 
them brown or black. Head and body, 380-400 mm. ; tail, 80-90 mm. 
(Thomas, 1888, pp. 245-246.) 

Its known range includes Tasmania and Victoria. In the former 
state "the bandicoots are very numerous everywhere; they . . . 
live principally on roots. I knew one gentleman's entire collection 
of Cape bulbs, principally Babianae, eaten by them, and I suffered 
considerably myself, having lost some entire species of bulbs 
through these animals." (Gunn, 1838, pp. 102-103.) Gray (in Gunn, 
1838, p. 108) records insect remains found in the stomach of one 

This species is to be met with throughout Tasmania, but it ap- 
pears to be less commonly and evenly distributed than the Short- 
nosed Bandicoot (Lord, 1928, p. 20) . 

"At one time distributed through western and central Victoria, 
this species is now restricted to a single locality near Hamilton and 
its numbers are few" (C. W. Brazenor, in litt., March 3, 1937). 

Western Barred Bandicoot; Marl 


Perameles myoswos Wagner, Archiv fur Naturg. (Wiegmann), 7th yr., vol. 1, 
p. 293, 1841. (The type locality, not stated in the original description, is 
Swan River, according to Glauert (1933, p. 23), or King George's Sound, 
West Australia, according to Iredale and Troughton (1934, p. 19).) 

FIGS.: Schreber, Saugthiere, suppl. 3, pi. 155 Ad, 1842; Gould, 1845, vol. 1, 
pi. 10. 

"No specimens have reached the [Perth] Museum since 1900. It 
is therefore assumed that the animal is extinct." (Glauert, 1933, 
p. 23.) 

Above mixed with blackish and yellowish brown; below dirty 
yellowish white; ears pale dusky, with a rusty-red spot at external 
base; a dark band extending across sides in front of thighs; feet 
whitish; tail scaly, short-haired, dusky above, dirty white below. 
Head and body, 11 inches; tail, 3 inches. (Wagner, 1841, pp. 293- 

The former range was the southwestern portion of Western 
Australia. According to Gould (1863, vol. 1, p. 14), it "inhabits 
the whole line of coast of the Swan River colony, but, so far as I 
can learn, is not found to the westward of the Darling range of 
hills." He adds that "its food consists of insects, seeds, and grain." 



"Apparently not plentiful in the South-west, although described 
by natives as being fairly numerous in the Salt River district. A 
species of Bandicoot, probably this species, is said to have formerly 
extended as far north on the mainland as Sharks Bay." Specimens 
are recorded from the vicinity of Pin jelly and Kojonup. (Short- 
ridge, 1910, pp. 833-834; map, p. 835.) 

FIG. 6. Western Barred Bandicoot (Perameles myosura myosura}. 
After Gould, 1845. 

South Australian Barred Bandicoot 


Perameles myosura notina Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 10, 
p. 144, 1922. ("Plains near the head of the St. Vincent Gulf," South 

FIGS.: Jones, 1924, p. 147, fig. 102; Le Souef and Burrell, 1926, fig. 83. 

This subspecies is apparently extinct in southeastern South 
Australia but is "probably holding its own in the semi-arid Nullarbor 
Plain" in the west (A. S. Le Souef, in Hit., February 15, 1937) . 

It closely resembles P. fasciata in coloration, with three distinct 
black bands on the hind quarters; skull with more slender muzzle 
and smaller teeth than in P. fasciata. Head and body, 280 mm.; 
tail, 90 mm. (Thomas, 1922, p. 144.) 

Formerly it seems to have ranged across the entire east-west 
extent of South Australia, in the more southern parts. According to 
Jones (1924, pp. 149-150), "this beautiful little Bandicoot had at 
one time a fairly wide distribution in this State. In addition to the 
animals from the head of St. Vincent Gulf, are others from the 
River Murray in South Australia, and from Adelaide itself. As 
far as can be ascertained it has now disappeared from all these 
localities, and remains only in the wastes of the western portion 
of the Centre. . 


"Barred Bandicoots become very tame and familiar in captivity, 
but . . . they are desperately pugnacious among themselves. On 
one occasion eight live specimens were sent from Ooldea. All eight 
were dead . . . when they arrived in Adelaide. . . . But among the 
corpses were four pouch young, which were uninjured. ... In the 
end they all recovered." A female from this lot eventually bred 
freely in captivity. Two young were generally found in a litter. 

Rabbit-eared Bandicoot; Rabbit-bandicoot; Bilby; Dalgite; 



Perameles Lagotis Reid, Proc. Zool. Soc. London 1836, p. 129, 1837. ("In 

Australia Occidentali et in Terra Van Diemen." Thomas (1888, p. 225) 

lists the type specimen from "Swan R., W. A.") 
FIGS.: Waterhouse, 1841, pi. 12, and 1846, vol. 1, pi. 13, fig. 1; Gould, 1845, 

vol. 1, pi. 7; Le Souef and Burrell, 1926, fig. 79; Pocock, 1937, p. 617, fig. 


The several subspecies of this Australian animal (lagotis, cam- 
brica, grandis, interjecta, nigripes, and sagitta) seem to be more or 
less seriously reduced in numbers; one is apparently on the verge 
of extinction, and another quite extinct. They will be treated in turn. 

Concerning the group as a whole, Jones says (1924, pp. 164-167) : 

By the early colonists the Bilby was not only regarded as an animal against 
which the methods of the exterminator need not be employed; it was even 
accorded a certain amount of protection, and was at times kept as a pet 
about the house. The tolerance with which it was regarded by people whose 
hands may be justly said to have been against all animals was due to the 
fact that it was recognised that, in the destruction of mice and insects, it 
played an extremely useful part. Unfortunately this regard for the Bilby 
seems to have been forgotten by a later generation, and in more recent days 
but little mercy has been shown to them by any section of the community. . . . 

The reason for the rapid decrease in numbers of the Bilby is not quite 
obvious. Certainly these useful animals have been ruthlessly slaughtered in 
all districts within reach of the more settled areas. Their pelts have been 
marketed in the skin sales in Adelaide in very large numbers; and they have 
been more wantonly killed for "sport." Large numbers have been killed or 
maimed in steel traps set for rabbits, and possibly many have fallen victims 
to poison baits. As with all the more defenceless marsupials, the introduced 
fox has probably played its sinister part. But in the Centre, where the fox 
is still absent, or rare, and where the Bilby is but little molested by man, it 
seems that some other factor must be invoked; and this is probably the 
extraordinary abundance of rabbits, and the consequent struggle for breeding 
burrows. There is certainly no part of this State [South Australia] where the 
Bilby is not a rapidly disappearing animal. 

Troughton remarks (1932, p. 221) : "According to Wood Jones, 
one or two constitutes the usual litter [in members of this genus], 
although there are eight teats, and it seems possible that a reduced 


rate of breeding, in the less hospitable regions to which settlement 
is forcing them, where the cunning introduced enemy [the fox] 
probably now abounds, must ultimately lead to the extinction of this 
harmless, picturesque, and pest-destroying marsupial." 

In the typical subspecies, M. I. lagotis, the general color is gray; 
head, neck, and back washed with chestnut; sides of body and base 
of tail pale chestnut; ears long, broad, ovate; outer, upper surface's 
of limbs grayish; under parts white; middle part of tail black; 
terminal part white, with a crest of stiff hairs. Head and body, 18J 
inches; tail, 10 inches. (Reid, 1837, pp. 129-130.) This is a large 
race, with a long, silky coat; the black portion of the tail usually 
longer than, or equaling, the crested white part (Troughton, 1932, 
p. 227). 

Its range includes south and central Western Australia and 
extends to Central Australia. 

Gould (1863, vol. 1, p. 11) considers it "tolerably abundant 
over the whole extent of the grassy districts of the interior of the 
Swan River colony." By retreating into its deep, long burrows, "it 
frequently eludes the pursuit of the natives, who hunt it for the 
sake of its flesh." He speaks of its flesh as "sweet and delicate," 
resembling "that of the rabbit." 

Its status in Western Australia is summed up by Shortridge 
(1910, pp. 832-833; map, p. 832) : 

"Although widely distributed throughout the South- West (except 
near the coast) , North-West, and Centre, it has within recent years 
become extremely rare in the far interior. Most plentiful in the 
inland districts of the South-West, rather frequently caught in traps 
set for rabbits along the rabbit-proof fence. In the dry North- 
Western and South-Eastern divisions, where it is rare, it extends to 
the coast." 

The same author states (1907, pp. 770-771) that in the interior 
"it seems to have almost left parts of the country where it was onca 
well known perhaps on account of the succession of droughts in- 
land of late years." 

Troughton (1932, p. 227) mentions specimens from Gracefield, 
Coorigan, and Teuterden, Western Australia. 

Glauert (1933, p. 24) records it in Western Australia as "widely 
distributed . . . south of the Kimberley Division. The western limit 
seems to be the Darling Range, although the Museum has odd 
specimens from Perth and Upper Swan on the Coastal Plain. The 
animal occurs as far south as Cranbrook and Jerramungup, near 
the Stirling Range, and as far east as Gnawlbat, 126 degrees 15 
minutes east, 26 degrees 21 minutes south." He writes (in litt., 


March 17, 1937) that it is "affected by fumigation of rabbit burrows 
in agricultural areas." 

Finlayson (1930, p. 178; 1931, p. 161) records specimens from 
north of the Musgrave Ranges, in the extreme north of South 
Australia, and from two localities north of the Macdonnell Ranges 
in Central Australia. 

E. Le G. Troughton writes (in Hit., April 16, 1937) that it "may 
survive always in parts of the Centre, but should be given total 
protection in the south-west as its destruction of rats and mice 
far outweighs any slight damage it may do." 

New South Wales Bilby 


Macrotis lagotis cambrica Troughton, Australian Zool., vol. 7, pt. 3, p. 230, 
1932. ("Bathurst," New South Wales.) 

This eastern subspecies, extinct in Victoria and last recorded 
from New South Wales in 1912, apparently survives in uncertain 
numbers in southern Queensland. 

It is about equal in size to the large western subspecies (lagotis) ; 
the fur is shorter and more woolly ; upper parts more fuscous ; under 
parts yellowish. Head and body, 390-500 mm.; tail, 248-278 mm. 
(Troughton, 1932, p. 230.) 

According to Troughton (1932, p. 230; map, p. 231), it was 
"originally distributed over inland New South Wales from the 
Darling River (Bourke) in the west, across to near the Great 
Dividing Range in the east (Bathurst and Ghoulburn) , south to the 
Murray River and north to the Queensland border (Moree) ; prob- 
ably extending into southern Queensland." 

Since 1892, fifteen specimens reached the Museum, making in all at least 
twenty-two, of which the last was received from Moree in June, 1908; though 
several are not definitely localised, there is no doubt that the entire series 
came from within New South Wales. . . . Probably never very plentiful, 
the rabbit-bandicoot was apparently distributed fairly evenly west of the 
dividing range in the early days, and, unless mere coincidence, appears to 
have been more abundant in some years as three specimens reached the 
Museum in August, 1897, and again in 1903. There seems no doubt, however, 
that the local race has vanished from the more settled areas, and that . . . 
the Bilby is rapidly disappearing from New South Wales, or is at least faced 
with a precarious existence in more desert regions. I am not aware of pelts 
ever having been marketed to any extent in Sydney, . . . but no doubt num- 
bers have been killed in rabbit traps, and wantonly for so-called sport, while 
foxes must be contributing to the apparent annihilation within New South 

The last record of the Bilby's occurrence in New South Wales, so far as I 
am aware, is [that of] a pair under close observation in the rocky hills on 
the Wagga Experimental Farm for about five years prior to 1912, when 


they were unfortunately slaughtered by shooters from the town. (Troughton, 
1932, p. 220.) 

In Victoria this Bilby was always confined to the northwestern 
corner of the state, and the last record was in 1860 (C. W. Brazenor, 
in Hit., March 3, 1937) . 

Jones (1923a, p. 342) speaks of examining a living specimen from 
Queensland. Finlayson states (1934, p. 229) that it is apparently 
absent from the Dawson Valley, Queensland, but that it occurs at 
Epping in the Clermont district, 150 miles northwestward. 

Nalpa Bilby 


Macrotis lagotis grandis Troughton, Australian Zool., vol. 7, pt. 3, p. 229, 1932. 

("Nalpa, in the Lake Alexandrina District, south of Adelaide, South 

Fia.: Jones, 1923a, p. 333, fig. 352. 

.This subspecies is apparently extinct (Troughton, 1932, p. 230). 

It is the largest subspecies; its ear, however, is proportionately 
shorter than in the other subspecies. Head and body, 550 mm.; 
tail, 260 mm.; ear, 77 mm. 

It is known only from the "South-east of South Australia" (Ire- 
dale and Troughton, 1934, p. 20) . 

Before this form was distinguished from the typical lagotis, 
Jones (1924, pp. 156-157) wrote of it as follows: 

Thalacomys lagotis, though formerly abundant in South Australia, is now 
either extinct or on the verge of extinction. It was the familiar species of 
Bilby in the more fertile portions of South Australia only a comparatively 
short time ago. Not more than thirty years since it was usual for rabbit 
trappers, even in the immediate neighbourhood of Adelaide, to take more 
Bilbies of this type than rabbits in their traps. This race . . . apparently had 
its last South Australian stronghold at Nalpa and in the wide tract of 
country about Lake Alexandrina; but from Nalpa it has long since disappeared, 
and it seems most probable that the animal is now extinct in this State. 

Rawlinna Bilby 


Macrotis lagotis interjecta Troughton, Australian Zool., vol. 7, pt. 3, p. 227, 
1932. ("Rawlinna, Trans-Australian Railway, Western Australia.") 

This subspecies seems to be known definitely from only two 
specimens, both taken at the type locality. 

^Smaller than the western lagotis; fur shorter and more woolly; 
general color more drab-gray and less contrasting; under parts 
whitish; black portion of tail equaling, or shorter than, the white 


terminal portion. Head and body, 303-318 mm.; tail, 207-232 mm.; 
ear, 80-82 mm. (Troughton, 1932, pp. 227-228.) 

"It is possible that the Musgrave Ranges specimen recorded by 
Finlayson [as lagotis] is an aged female of this race" (Troughton, 
1932, p. 228) . 

Black-footed Bilby 


Thalacomys nigripes Jones, Rec. S. Australian Mus., vol. 2, no. 3, p. 347, 
figs. 358-360, 1923. ("Ooldea Soak," Trans-Australian Railway, South 

FIGS.: Jones, 1923o, p. 347, fig. 358, and 1924, p. 152, figs. 106, 107. 

"This animal is, so far, only known from the district round 
Ooldea Soak, on the railway from Port Augusta to Perth. In that 
district it appears to be by no means uncommon." (Jones, 1924, 
p. 163.) 

Smaller than M. I. lagotis but much like it in general color; 
distinguished from all other members of the genus by its black 
feet; under parts pure white; black portion of tail shorter than 
terminal white portion. Head and body, 365-390 mm.; tail, 200- 
220 mm.; ear, 105-110 mm. (Jones, 1923a ; pp. 347-350.) 

The half-dozen known specimens were all collected by abo- 
riginals (Jones, 1923a, p. 349) . 

"On the Nullarbor Plain, in the state of South Australia, occa- 
sional holes would be met with, the animals numbering, in 1921, 
about 21 to the square mile" (Le Souef and Burrell, 1926, p. 299). 
Although these authors add that the form was probably sagitta, 
there would seem to be equal or greater likelihood of its being 
nigripes, since the type locality of the latter borders on the Null- 
arbor Plain. 

Barcoo Bilby 


Thalacomys sagitta Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 16, p. 426, 1905. 
("Killalpanima [= Killalpaninna], east of Lake Eyre," South Australia.) 

Although still considered "by no means uncommon" (Finlayson, 
1935c, p. 233), this Bilby, like other members of the genus, is 
probably seriously menaced by the predatory fox and the com- 
peting rabbit. 

This is smaller than any subspecies except interjecta, and a 
little paler than lagotis. Head and body, 316-385 mm.; tail, 215- 
245 mm.; ear, 79-84 mm. (Thomas, 19056, p. 426; Troughton, 
1932, p. 229; Finlayson, 1935c, pp. 234-236.) 


The Barcoo Bilby has been recorded in northeastern South 
Australia, from Miller's Creek and Coward Springs, southwest of 
Lake Eyre, to the Goyder's Lagoon area toward the northeastern 
corner of the state. It also seems to range northward to the region 
about Charlotte Waters, Central Australia. (Jones, 1923a, p. 344; 
Finlayson, 1935c, p. 233.) 

Jones writes (1924, p. 160) : "This Bilby is a northern form 
living in the region of the great drainage system of Lake Eyre. It 
is probable that it is still fairly abundant in those portions of this 
legion where foxes have not yet become plentiful, and where it 
can still compete with rabbits for nesting burrows." 

On this subject Troughton says (1932, p. 221) : "My own 
experience when collecting in the very dry country about Farina 
[south of Lake Eyre], South Australia, in 1919, was that foxes 
were very numerous and already tending towards a small lean 
desert type capable of entering the larger rabbit burrows without 
difficulty, and doubtless those of the Bilbies as well." 

Six specimens obtained in the Goyder's Lagoon area about 1932 
were, according to Finlayson (1935c, p. 233), the first ones to be 
examined in the flesh since the type specimen was taken in 1903. 

Certain notes on "Peragale lagotis" from the Charlotte Waters 
region of Central Australia, published by Spencer (1896, p. 17, and 
1897, p. 9) before sagitta was recognized, actually refer, it seems, 
to the latter form (c/. Troughton, 1932, p. 233). "This is not 
uncommon, judging by the number of tails used by the natives 
as ornaments. They tie the white terminal tufts together in bundles 
of from twelve to twenty." The animal occupies the inner end 
of its burrow, and the natives secure it by digging it out. 

White-tailed Bilby; White-tailed Rabbit-bandicoot 


Peragale leucura Thomas, Ann. Mag. Nat. Hist., ser. 5, vol. 19, p. 397, 1887. 

("Exact locality . . . not . . . recorded.") 
SYNONYM: Thalacomys minor miselius Finlayson (1932). 
FIGS.: Thomas, 1888, pi. 2; Finlayson, 1935, pi. opp. p. 63. 

According to Finlayson (1935c, p. 232), the probability of the 
identity of M. I. leucura and T. m. miselius is very great, and the 
latter name is here considered, at least provisionally, as a synonym. 

Proportions and fur of leucura as in M. lagotis; general color 
pale yellowish fawn; under parts pure white or yellowish white; 
limbs pure white; tail slender, wholly white-haired, with a terminal 
dorsal crest. Measurements of the very young type: head and 
body, 142 mm.; tail, 116 mm. (Thomas, 1887, pp. 397-398.) In 
miselius the central two-fifths of the tail has a median dorsal line 


of pale slate, bordered by fawn; head and body, 250 mm.; tail, 
155 mm.; ear, 72 mm. (Finlayson, 1932, pp. 168-169). 

The type of leucura, from an unknown Australian locality, was 
described in 1887. A second specimen was taken at Mungerani, 
east of Lake Eyre, in 1924. The 12 specimens on which the name 
miselius was founded were taken in 1931 near Cooncherie on the 
lower Diamantina River, in northeastern South Australia, at about 
latitude 26 32'. In this area the animal was plentiful (Finlayson, 
1935c ; p. 227). It appears to be known, however, from a total of 
only 14 specimens. 

A Wonkonguroo boy, who obtained most of the specimens near 
Cooncherie, was adept at locating the burrows in sand hills, 
although the entrances were blocked with loose sand. The animal 
is evidently used as food by the natives. (Finlayson, 1935c, p. 227.) 

"It now appears . . . that the ... composite species [M. I. 
leucura and M. I. minor] has a wide central distribution in which 
it may survive indefinitely, though the advent of the fox and 
rabbit are considered by Professor Wood Jones to have already 
exercised a marked influence on sub-desert populations, in asso- 
ciation with prolonged dry seasons" (E. Le G. Troughton, in litt., 
April 16, 1937) . 

Lesser Bilby; Lesser Rabbit-bandicoot 


Peragale minor Spencer, Proc. Royal Soc. Victoria, n. s., vol. 9, p. 6, pi. 2, 
figs. 1-4, 1897. ("Sand-hills about forty miles to the north-east of 
Charlotte Waters," Central Australia.) 

This animal seems to be definitely known only from a small 
series taken at the type locality in Central Australia more than 
45 years ago. 

Fur long and silky; general color fawn-gray; chin and inner 
side of forelimbs white; rest of limbs and under parts gray; feet 
white above; basal two-thirds of tail dark above; final third 
white, crested; sides and ventral surface of tail white. Head and 
body, 200-270 mm.; tail, 118-160 mm.; ear, 68-92 mm. (Spencer, 
1897, pp. 6-7.) 

"The 'Urpila' (P. minor) during the winter months lies within a 
foot or so of the entrance of his [burrow]. . . . This peculiarity 
is taken advantage of by the natives who jump on the surface of 
the ground behind the 'Urpila' breaking it in and so cutting off 
his retreat to the inner chamber. He is thus compelled to rush 
out through the entrance where a native is waiting to give him 
his quietus." (Byrne, in Spencer, 1897, p. 9.) 


The fox and the rabbit have probably been decisive factors in 
the depletion or disappearance of this animal. 

Eastern Pig-footed Bandicoot 


Perameles ecaudatus Ogilby, Proc. Zool. Soc. London 1838, p. 25, 1838. (Left 
(south) bank of the Murray River, near the junction with the Murrum- 
bidgee River, in Victoria. Not New South Wales, as stated by Iredale 
and Troughton (1934, p. 21). Cj. Mitchell, 1838, vol. 2, p. 131.) 

FIGS.: Gould, 1845, vol. 1, pi. 6 (central fig.); Jones, 1924, p. 167, fig. 124. 

This unique little animal has apparently vanished from eastern 
Australia and southern South Australia; possibly it maintains a 
slight foothold (as one subspecies or the other) in Central Australia. 

Ears long, elliptical, and nearly naked; muzzle much attenu- 
ated; body about the size of a small rabbit, and the fur very 
much of the same quality and color as in that animal; two toes 
on forefeet, similar to those of a pig; tail [accidentally] wanting 
(Ogilby, 1838, pp. 25-26). General color coarsely grizzled gray, 
with a tinge of fawn; under parts white; limbs long and slender; 
tail black above, gray below and on sides. Head and body, 250 
mm.; tail, about 100 mm. (Thomas, 1888, pp. 251-252.) 

The former range included the interior parts of Queensland, New 
South Wales, and Victoria; also South Australia. The systematic 
status of the Central Australian animal is apparently not settled, 
but in coloration it is said by Spencer (1896, p. 17) to resemble 
the western subspecies, C. e. occidentalis. 

"The quaint and singularly gentle Pig-footed Bandicoot which 
had been discovered by Mitchell in 1836 was reported by Krefft 
twenty years later as exceedingly rare and disappearing as fast as 
the native population" (Troughton, 1932, p. 188). This was due 
to the increase of cattle and sheep (Lydekker, 1894, p. 148) . 

The species is recorded from western Queensland by Longman 
(1930, p. 64). 

There were a few records from extreme northwestern Victoria 
(the last one in 1857), and the animal is now extinct in that state 
(C. W. Brazenor, in litt., March 3, 1937). 

Jones writes (1924, p. 171) concerning the species in South 
Australia : 

Specimens in the South Australian Museum come from Cooper's Creek, 
from near Ooldea, and from the Gawler Ranges. Probably it still lives in 
the neighbourhood of Ooldea, but specimens have not been met with in 
that district for some years. ... In 1920 one was killed between Miller's 
Creek and Coward Springs to the south and west of Lake Eyre. . . . Although 
its distribution in the Centre is wide, it has always been a very rare animal, 
and now must be regarded as a disappearing one. . . . 


Pig-footed Bandicoots are said . . . when chased by dogs, to seek the 
shelter of hollow logs or hollow trees. In the districts to which they are now 
confined they would be hard put to find a log, let alone a tree .... Once 
open country of this type has been invaded by the fox, the fate of Choeropus 
is sealed. . . . The name by which it is known to the Kukata blacks is 
Wilalya, and they regard it as an animal which has always been rare and 
which is now extinct in their country. 

Reporting on the Horn Expedition to Central Australia, Spencer 
says (1896, pp. 17-18) : 

"At the present time this is one of the most difficult of the smaller 
marsupials to secure. . . . During the expedition we were unable 
to secure a single specimen. On a subsequent visit to Charlotte 
Waters I was fortunate enough to obtain one secured by the 
blacks. . . . 

"There is no doubt but that the range of the animal extends 
widely over the central area. In the Adelaide Museum is a speci- 
men from Barrow Creek, which lies well within the tropics, and 
throughout the whole of our expedition all the natives were well 
acquainted with it. ... It ... is evidently rapidly becoming ex- 
tinct, except perhaps in the more central districts." 

"Said to still have a wide but sparse distribution in the central 
region, there has been little proof of late, and its terrestrial, non- 
burrowing, specialized habits and frail constitution render its ulti- 
mate extinction certain" (E. Le G. Troughton, in litt., April 16, 

Western Pig-footed Bandicoot 


[Choeropus] occidentalis Gould, Mamm. Australia, vol. 1, p. 10, pi. 6, 
1845. ("The interior" of "Western Australia"; type locality shown by 
Thomas (1888, p. 252) to be "Boorda, Kirltana, W. A.") 

FIGS.: Gould, 1845, vol. 1, pi. 6 (right and left figs.) ; Waterhouse, 1846, vol. 1, 
pi. 13, fig. 2. 

This animal is extinct, at least in Western Australia (L. Glauert, 
in litt., March 17, 1937). The form that once occurred in Central 
Australia (see discussion under C. e. ecaudatus) does not seem to 
have been recorded for some years and may have suffered the 
same fate. 

The western subspecies differs from the eastern one chiefly in its 
orange-brown rather than gray coloration. 

Gould states (1863, vol. 1, p. 10) that Gilbert sent two specimens 
from Western Australia and that the animal is confined to the 
interior. According to Waterhouse (1846, vol. 1, p. 392), one of the 
specimens came from the Swan River district. 

"I was not able to find out anything definite about the dis- 


tribution of this species in Western Australia. It is evidently very 
rare. The specimen obtained by Gilbert in 1843 seems to have been 
the only one ever secured in this State." (Shortridge, 1910, p. 835; 
map, p. 836.) 

"Mr. A. Le Souef states in a letter 2/12/1927 that he has seen a 
dried skin at Rawlinna. This is the only recent record known to 
me." (Glauert, 1933, p. 24.) 

Family PHALANGERIDAE : Phalangers, etc. 

This family consists of approximately 14 Recent genera and 110 
forms. Its range extends from Tasmania and Australia to New 
Guinea and the Admiralty and Solomon Islands on the north and 
to Celebes and Timor on the west. Three Australian species are 
discussed herein. 

Honey Mouse; Honey Possum; Long-snouted Pouched Mouse 


Tarsipes Spenserae J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 9, p. 40, 1842. 

("King George's Sound," Western Australia.) 
FIGS.: Gould, 1845, vol. 1, pi. 5; Waterhouse, 1846, vol. 1, pi. 11, fig. 1; 

Cabrera, 1919, pi. 13, fig. 5; Troughton, 1923, pi. 23, and p. 152, fig.; 

Troughton, 1924, pp. 128, 129, figs. 

This rare, local, and unique little animal of Western Australia is 
"becoming rarer" (L. Glauert, in Hit., March 17, 1937) . 

Body mouselike; head elongate, tapering; general color blackish 
gray ; back with a median black streak, bordered by a brown stripe 
on each side; under parts pale bay; tail elongate, tapering, short- 
haired, scaly. Head and body, 3^ inches; tail, 3 inches. (J. E. Gray, 
1842, p. 40.) Tongue slender, protrusile, and brushlike, specialized 
for thrusting into flowers for nectar. Head and body, 71 (male) to 
86 mm. (female) ; tail, 95 (male) to 101 mm. (female). (Troughton, 
1923, pp. 153-154.) 

The range, according to Glauert (1933, p. 25), is "South-Western 
Australia from the Irwin River south of Geraldton to the south 
coast as far east as Esperance. 

"Usually more or less coastal, but has been found along the 
Great Southern as far north as Wagin, and at Nyabing east of 
Katanning. The animal still occurs close to Perth in suitable 
localities. . . . King George's Sound ... is still the headquarters 
of the species." 

Gould (1863, vol. 1, p. 9) recorded it "from Swan River to King 
George's Sound, but from its rarity and the difficulty with which it 
is procured, notwithstanding the high rewards I offered, the natives 
only brought me four specimens." 


Shortridge (1910, p. 826; map, p. 827) records eight specimens 
from Albany. "The small marsupial mice are very difficult to secure 
on account of their rarity, and their nocturnal, arboreal, and to a 
great extent insectivorous habits, being known chiefly from cats 
killing and bringing them into houses." 

Troughton, who has contributed most of the recent information 
on the species, writes (1924, pp. 127-132) : 

Alas, as settlement increases, the time seems near when there may be no 
living representatives of these unique creatures to occupy the queer niche 
which the process of evolution has fashioned for them within its fabric. . . . 

Failure [to secure specimens on a collecting trip near Albany in 1922] 
was not surprising, all accounts confirming Mr. Morgan's statement that the 
mice visit areas periodically according to the flowery food supply, and that 
they are but rarely seen except when brought in by cats as trophies of the 
chase. . . . 

A few months after my return . . . , the Honey Mice visited Mr. Morgan's 
home once more and he has since sent over twenty adult mice to the Museum, 
all of which were caught by his cat. . . . 

As they are dependent upon the native flowers, the advance of settle- 
ment with its periodical burning off, and the introduction of cats and other 
enemies, in addition to native ones, must seriously threaten the future of 
these marsupials .... 

It is reassuring to hear from Mr. Glauert that the mice are still fairly 
plentiful over an area of about 12,000 square miles, and that at the end of 
1923 the West Australian Government was about to proclaim the Stirling 
Ranges a sanctuary for the native fauna. Let us hope that these ranges may 
prove a veritable stronghold for the Honey Mice, and that the flowers may 
not miss their spring-cleaning from the brushy tongues for many generations 
to come. 

Troughton also says (1923, p. 155) : "Tarsipes is dependent upon 
the flowers and foliage of its native districts, and as paddocks have 
to be burnt off about every third year, the tiny marsupials are 
literally hunted from paddock to post and prevented from settling 
in any one area. Before the rapid advance of Western Australia's 
settlement schemes, . . . fire and other enemies will send the Honey 
Mice to join their fossil forbears in comparative oblivion, leaving 
them represented only by a few museum skins and stray skeletons." 

Leadbeater's Opossum 


Gymnobelideus Leadbeaten M'Coy, Ann. Mag. Nat. Hist., ser. 3, vol. 20, p. 287, 

pi. 6, 1867. ("Banks of the Bass River, in Victoria.") 
FIGS.: M'Coy, 1867, pi. 6; McCoy, 1883, pi. 91; Lucas and Le Souef, 1909 ; 

p. 107, fig.; Brazenor, 1932, pi. 6. 

This rare species, the only known member of its genus, was based 
upon two specimens collected in 1867 along the Bass River, South 
Gippsland, Victoria. In 1900 another specimen, reputed to have 


come from the same locality, was secured from a Melbourne dealer. 
In 1909 the fourth known specimen was presented to the National 
Museum of Victoria by A. G. Wilson; it came from Mount Wills in 
East Gippsland, some 160 miles from the Bass River habitat. A fifth 
specimen was presented to the same institution by F. V. Mason, 
who had taken it many years previously at the edge of the Koo- 

FIG. 7. Leadbeater's Opossum (Gymnobelideus leadbeateri) 

Wee-Rup Swamp, about 3 miles due south from Tynong Railway 
Station and only a few miles from Bass River. (Brazenor, 1932, 
pp. 106, 108.) 

Practically our entire knowledge of the species is based upon these 
five specimens, which are in the National Museum of Victoria. 

Its general appearance is much like that of the Sugar Glider 
(Petaurus breviceps) , but it has no flying membrane. Color above 
brownish gray to fawn-gray, with a dark brown to black median 
stripe from head to sacrum; dark patches above and below ear and 
about eye; ears large, nearly naked toward tip; chin and throat 
dull buff; rest of ventral surface light yellowish gray; tail long, 
bushy, colored like body (in one specimen the terminal half is 
black) . Head and body, 169-200 mm.; tail, 168-203 mm. (Brazenor, 
1932, pp. 106-108.) 


"There is much virgin scrubland in Gippsland in which the small 
creature could survive. It is nocturnal in habits, and its general 
resemblance to Petaurus breviceps is close enough to make its recog- 
nition by evening light very difficult. When these facts are con- 
sidered, the possibility of its survival is greater than might at first 
be realised, and it is probable that a systematic search would re- 
establish this small creature among the living animals of Victoria." 
(Brazenor, 1932, p. 109.) More recently Mr. Brazenor has come 
to the conclusion (in litt., March 3, 1937) that it is "probably 

"The risk of extermination for small unexploitable opossum forms 
of restricted range is indicated by the fate of Leadbeater's opossum 
. . . , an important phalangerid link, originally restricted to a small 
area of Victoria where denudation of its limited scrub habitat 
has apparently led to the animals' extinction. This unique mar- 
supial is represented by barely a dozen specimens in state museums." 
(Troughton, 1938, p. 408.) 

Presumably Leadbeater's Opossum has been subject 'to attack by 
the Domestic Cat and perhaps other introduced enemies. It is also 
quite possible that many specimens have fallen victims to opossum 
trappers who did not differentiate them or at any rate did not 
realize their exceptional value. 

Some of the numerous government reserves that have been 
established in Victoria might provide sanctuary for this unique 
animal if it were still extant. 

Western Ringtail; Western Ring-tailed Opossum 


Pseudochirus occidentalis Thomas, Cat. Marsupialia and Monotremata Brit. 
Mus., p. 174, 1888. ("King George's Sound, W. A.") 

This animal, confined to the extreme southwest of Western 
Australia, is feared to be on the verge of extinction. 

Color above deep smoky gray; limbs like back, but hands and 
feet darker; under parts white; basal part of tail dark brown, 
terminal two-fifths white, naked part below tip smooth. Head and 
body, 335 mm.; tail, 310 mm. (Thomas, 1888, pp. 174-175.) 

Nearly a hundred years ago Gould's collector, John Gilbert, 
obtained specimens at Perth, Swan River, and King George's Sound. 
More recently Shortridge (1910, pp. 827-829) collected 22 specimens 
at Margaret River and Busselton, remarking that it is "chiefly 
confined to the banks of rivers and swamps in the South-West; 
local, and apparently disappearing in many places." Yet he con- 
sidered it "fairly plentiful near the Margaret River." These few 


records suggest that the animal was practically confined to the 
South-West Division of Western Australia, from Perth southward. 
(See map, Shortridge, 1910, p. 829.) 

Glauert (1933, p. 24) gives its range as "lower South- Western 
Australia in small isolated colonies, which suggest that the animal 
is on the verge of extinction through natural causes." 

No particular information is at hand concerning its enemies, aside 
from Gilbert's remark (in Gould, 1863, vol. 1, p. 25) : "It ... is 
often found in holes in the ground, . . . from which it is often 
hunted out by the Kangaroo dogs." 


The Koalas, consisting of a single genus and species, with three 
subspecies, are restricted to eastern Australia. All forms come 
within the scope of the present work. 

New South Wales Koala; Native Bear 


fypurus cinereus Goldfuss, in Schreber, Saugthiere, pis. 155 Aa, Ab, 1817; 

Isis (Oken), 1819, Heft 2, p. 271. ("The forests of New Holland, about 

50-60 English miles [southwest] from Port Jackson [Sydney]," New 

South Wales.) 
FIGS.: Waterhouse, 1841, pi. 31; Gould, 1854, vol. 1, pis. 13, 14; Lydekker, 

1894, pi. 10; Nat. Geog. Mag., vol. 70, no. 6, p. 715, right-hand fig, 1936; 

Pocock, 1937, p. 626, fig. (subsp.?). 

Once numerous in the timbered areas of New South Wales, the 
typical subspecies of this unique animal has been reduced almost to 
the verge of extinction, although many thousands of the Queensland 
subspecies (P. c. adustus) and perhaps a thousand of the Victorian 
subspecies (P. c. victor) still exist. 

The fur is dense and woolly; general color gray, either light or 
dark, sometimes mottled, with whitish patches on hind quarters; 
under parts, hands, and feet more or less whitish ; ears large, thickly 
haired; tail rudimentary. Head and body, 700-820 mm. (Le Souef 
and Burrell, 1926, pp. 291-292.) Auburn groin patches separated 
by a creamy-white median area (Troughton, 1935, p. 139). 

The Koala feeds almost entirely on the foliage of a few trees 
of the genus Eucalyptus: E. viminalis, E. melliodora, E. rostrata, 
E. microcorys, and E. maculata (Sutton, 1934, p. 78). Thus the 
ranges of the three subspecies are pretty definitely restricted to 
those areas in which /some or all of these eucalypts occur. The 
species as a whole formerly ranged from extreme southeastern 
South Australia through Victoria and the eastern half of New 
South Wales into Queensland (see map, Victorian Nat., vol. 51, 


no. 3, p. 80, 1934). While the exact geographical limits of the 
several subspecies have not been fully determined, we may pro- 
visionally consider the range of cinereus to be New South Wales; 
of adustus, Queensland; and of victor, Victoria and southeastern 
South Australia. 

In New South Wales Gould (1863, vol. 1, pp. 18-19) considered 
the animal "nowhere very abundant" but most numerous "in the 
brushes which skirt the sea side of the mountain-ranges between 
the district of Illawarra and the River Clarence." He recorded it 
also "among the cedar brushes of the mountain ranges of the interior, 
particularly those bordering the well-known Liverpool Plains." He 
prophesied that it "is certain to become gradually more scarce, and 
to be ultimately extirpated." 

"Though at one time extremely numerous, the koala is now, over 
the greater part of its range, very scarce. This is largely due to 
a disease which swept it off in millions in the years 1887-8-9, and 
from 1900 to 1903. This disease took the form of ophthalmia and 
periostitis of the skull. Bears are generally heavily infected with 
intestinal parasites." (Le Souef and Burrell, 1926, p. 292.) 

At Marrangaroo, County of Cook, N. S. W., "the native bear 
was quite common then [1884-5], but quite extinct there now" 
(Chisholm, 1923, p. 60) . On the Comboyne Plateau, N. S. W., it 
is "very rare here now and only inhabiting the Eucalypt timber at 
the edge of the Plateau" (Chisholm, 1925, p. 72). 

In the fox-free eastern coastal area "there are also a few Koalas, 
but these never get a chance, as the temptation to shoot or catch 
the defenceless little animals as they sit exposed on a bough, is 
more than the so-called sportsmen of the community can resist, 
and even in our National Parks they are destroyed" (Le Souef, 
1923, p. 110). 

Barry writes (1928, p. 163) of the Koala's status on Kuringai 
Chase, near Sydney: "Native Bears were also common here years 
ago, but now, as in most places in New South Wales, they are 
rarely seen." 

"The typical N. S. Wales animal has been reduced to a state 
verging upon extinction, in which the setting aside of adequate 
areas with assured supply of favoured eucalypt diet trees presents 
the only hope of survival. It is notable that any attempt to breed 
them in captivity is dependent for ultimate success on the provision 
of such reserves." (E. Le G. Troughton, in litt., April 16, 1937.) 

Lydekker remarks (1894, p. 80) in regard to the Koala's economic 

The flesh is considered a great delicacy by the natives, and is regarded as 
not unpalatable even by Europeans. Of its pursuit by the natives in the neigh- 
bourhood of Port Jackson, Colonel Patterson writes as follows: "The natives 


examine . . . the branches of the loftiest gum-trees, and upon discovering a 
Koala, they climb the tree in which it is seen with as much ease and 
expedition as a European would mount a tolerably high ladder. . . . They 
follow the animal to the extremity of a bough, and either kill it with the 
tomahawk or take it alive. . . ." 

The Koala must be an abundant animal, since from 10,000 to 30,000 skins 
are annually imported into London, while in 1889 the enormous total of 
300,000 was reached. The value of these skins now ranges, according to Poland, 
from five-pence to a shilling each; and they are mainly used in the manu- 
facture of those articles for which a cheap and durable fur is required. 

Concerning the Koala's decline, persecution, and need of protec- 
tion, Jones writes (1924, pp. 184-186) : 

It may be said to spend its whole life clinging to, and feeding upon, the 
great eucalyptus trees. In just so much as it is a perfected specialisation 
to its environment, so it is a slave to its environment. It has adapted itself to 
the gum tree, and has become dependent upon the gum tree. It must be 
regarded as an animal which has become phylogenetically senile as the out- 
come of complete specialisation .... 

Probably no animal has been so ruthlessly slaughtered in order to satisfy 
the demands of the fur trade. ... In the year 1908, no less than 57,933 Koala 
pelts passed through the markets of Sydney alone. That this deplorable 
slaughter still goes on is evidenced by the fact that in the two years 1920 
and 1921 Osborn and Anthony have ascertained that the huge total of 
205,679 Koalas were killed for the fur market. Since in the fur trade Koala 
pelts pass under the name of "Wombat," many people assume that the 
Native Bear has ceased to be persecuted. 

The complete extermination of the Native Bear would be a disgrace to 
Australia, and yet, from its dependence upon a particular diet and a par- 
ticular mode of life, its tenure of continued existence must always be regarded 
as precarious. . . . 

Horrible cruelties have been committed and recounted by those who have 
slaughtered them wholesale for the sake of their pelts. Indeed, one may say, 
on humanitarian grounds, that not only should the slaughter of the Koala for 
the fur trade be prohibited because the animal is eminently one to protect 
and not to exterminate, but it should be prohibited because, like the slaying 
of seals, it is the most brutalising occupation that a human being can 

Le Souef and Burrell say (1926, pp. 291-292): "The quaint 
koala . . . , perhaps, holds the affection of Australians more than 
any other of their wild animals a fact for which its innocent, 
babyish expression and quiet and inoffensive ways are largely re- 
sponsible. It has been portrayed in caricature and verse, and its 
hold on the public is used effectively by advertisers. . . . 

"The skin forms a thick, serviceable fur that will stand any 
amount of hard usage. Only the most callous of shooters, however, 
can bring themselves to shoot such a childlike animal." 

"In each of the States of Victoria, Queensland and New South 
Wales, the animal is protected by law" (Stead, 1934, p. 18). Im- 
portation into the United States of America was prohibited about 
1930-31 by the United States Government. "So long as the United 



States market remains closed there will be but little local incentive 

for destruction quite apart from any Australian protective laws" 

(Wild Life Preservation Society of Australia, 22d Ann. Kept., 1931) . 

Hobley calls attention (1934, p. 79) to private sanctuaries for 

FIG. 8. New South Wales Koala (Phascolarctos cinereus oinereus) 

the Koala established by Noel Burnet in the Pennant Hills, near 
Sydney, and by C. A. M. Reid at Lone Pine, near Brisbane. (The 
latter is presumably stocked with the Queensland subspecies.) He 

"Great credit is due to the founders of these sanctuaries who 
have been public-spirited enough to secure the safety of a number 
of these creatures without any Government support. . . . 


"The Wild Life Preservation Society of Australia is also wisely 
working away with the object of establishing Koala Colonies in 
such places as the Jenolan Caves Sanctuary, Lindfield Park, and 
Davidson Park in New South Wales the State authorities must, 
however, be persuaded to guarantee some security for the animals 
established therein." 

The cause of the Koala is eloquently pleaded by Troughton 
(1932a, p. 192) : 

"The Koala ... is utterly harmless everywhere, and what a 
delight it would be for both young and old if they were plentiful 
enough to haunt the suburbs and homesteads as possums often do. 
They seek only the freedom of the trees, and if the continued 
slaughter of such innocents leads to their extermination, it must 
inevitably appear to later generations as an indictment of the 
cultural degradation of our time." 

Queensland Koala 


Phascolarctos cinereus adustus Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 11, 
p. 246, 1923. ("0 Bil Bil, near Mimdubbera," Eidsvold, South Queensland.) 

FIGS.: Faulkner, Australian Zool., vol. 3, pt. 3, pi. 16, 1923; Le Souef and 
Burrell, 1926, fig. 77. 

Up to 15 or 20 years ago, the Queensland Koala must have 
numbered well over a million individuals; quite possibly there 
were several millions. But disease and more especially the fur 
trade have reduced it to a remnant of its former numbers. 

It is smaller than P. c. cinereus; fur shorter; anterior back suf- 
fused with dull rufous or tawny; ears far less thickly hairy, the 
inner surface almost naked; under parts lighter; the prominent 
groin patches rather browner and less rufous. Head and body, 
600 mm. (Thomas, 1923, p. 246.) 

In former times it seems to have ranged over practically all 
the more southerly and easterly parts of Queensland, north to 
Inkerman (lat. 19 30') and west to the Diamantina and Cooper 
River basins (about long. 143). (See map, Victorian Nat., vol. 51, 
no. 3, p. 80, 1934.) 

"The Queensland Minister for Agriculture has said that in 
1919-1930 no fewer than ... a million native bears were slaugh- 
tered in Queensland. If this slaughter continues these poor animals 
will be exterminated." (Gregory, 1921, p. 65.) 

"Koalas . . . are now getting numerous again in Southern Queens- 
land" (Le Souef, 1923, p. 109). 

"In 1924, the colossal total of over two million skins of the 
Koala or Native Bear were exported and mainly sold under the 


name of 'wombat' to mask the wholesale slaughter. In the Queens- 
land open season of 1927, approximately 600,000 Koala were mas- 
sacred by 10,000 licensed trappers." (Troughton, 1932, p. 193.) 

"In Queensland Native Bears are still to be found in fair numbers, 
and no doubt the Queensland Government was influenced by this 
fact when it removed the protection which the animals have enjoyed 
since 1919. But it is certain that even in one month their numbers 
will be seriously depleted. . . . Fur and skin brokers in Brisbane 
considered that before the season closed 300,000 skins would have 
been disposed of. It is doubtful whether this estimated total will 
have been reached, but it has to be remembered that many young 
will perish when deprived of the parental care of their mothers, 
which carry the little ones 'pick-a-back' from June until towards 
the close of the year." (Anonymous, 1927, p. 112.) 

Stead (1934, pp. 16-17) writes: 

Only in a few places in Queensland are large numbers of the Koala to be 
found, but only the most careful protection by the Government and by the 
Australian people will prevent them from being exterminated in these places. 
. . . Telling of the terrible destruction which has gone on in Queensland . . . 
makes a very unhappy story, and makes one rather ashamed to think that 
his own people should so cruelly destroy one of the most fascinating, harmless 
and most interesting living things in the whole of the world of Nature. . . . 

Very few people have any idea of the immense number of these harmless 
animals killed in the one State of Queensland in only a few years before the 
present season of protection was introduced. In 1927, about 600,000 were 
killed during one month's open season (August), and, for the whole year, 
including a so-called close season, not less than one million were slain. 
Altogether, several millions of the poor little Koalas were killed in a space 
of a few years in Queensland, until a great wave of public indignation put a 
stop to it for the time being. 

"The tenure of the koala in the Dawson Valley [Queensland] 
seems to have been a 'waning one for many years, and the last open 
season reduced it to such an extent that it is now a rare animal in 
many parts of the valley where it was formerly very plentiful. The 
process has been hastened, too, in some places, by an epidemic, and 
on Coomooboolaroo in the summer of 1929 several were seen in 
comatose condition at the base of feeding trees. The single example 
in this condition which was examined closely was an aged male, 
and though emaciated was not heavily infested with endoparasites, 
nor obviously diseased organically. . . . 

"It was observed .and collected at Thangool on the Cariboe, at 
Coomooboolaroo, and near Mount Hedlow, on the Fitzroy." (Fin- 
layson, 1934, p. 220.) 

The animal now has complete legal protection in Queensland 
(Stead, 1934, p. 18) . 

"Perpetual universal protection is essential to its ultimate sur- 


vival, not alone in New South Wales and Victoria where exploita- 
tion, denudation of habitat, and disease have brought this unique 
animal to the verge of extinction. It is in Queensland especially, 
where coastal forests of the south-east provide the last stronghold 
and hope of survival, that total protection should never again be 
withdrawn." (Troughton, 1938, p. 408.) 

Victorian Koala 


Phascolarctos cinereus victor Troughton, Australian Nat., vol. 9, pt. 6, p. 139, 

1935. ("French Island," Western Port Bay, Victoria.) 
FIGS.: Victorian Nat., vol. 51, no. 3, pis. 11-15, 1934. 

Once very common over most of Victoria and in southeastern 
South Australia, this subspecies had become reduced by 1934 to 
about 1,000 individuals in Victoria. 

Body more robust than in P. c. cinereus; coat longer, sparser, 
and hairier, especially on rump and ears; coloration richer, de- 
cidedly brown; ears brownish outside, white inside; auburn groin 
patches extending across the inguinal region; belly brown (Trough- 
ton, 1935, p. 139) . White area on throat and chest frequently pro- 
longed to nape, forming a complete collar. Head and body: three 
males, 800-830 mm.; one female, 730 mm. (Finlayson, 19356, pp. 

The Koala's status in Victoria is thus reviewed by Lewis (1934, 
pp. 73-74) : 

There is very good evidence that forty or fifty years ago "Native Bears" 
were exceedingly common over almost the whole of Victoria. Now the species 
is almost extinct on the mainland, a very few Koalas surviving in the 
Inverloch district and in South Gippsland around Welshpool, Toora, Foster, 
etc. Others are living and, I am glad to say, thriving on the islands in 
Western Port Bay. I estimate that there are now not more than 1,000 Koalas 
in this State. 

On the mainland of Victoria, I feel certain, the Koala is doomed to early 
extinction, and will never be re-established, excepting perhaps in some reserves 
which may be specially set apart for its protection and conservation, such as 
the Badger Creek Sanctuary, near Healesville. . . . 

From inquiries I have made among well informed people, it appears that 
the favourite "sport" of the young men and boys of thirty or forty years ago 
was shooting Native Bears. Their ideas of "sport" must have been very 
primitive, because no more inoffensive and easily-destroyed animal than the 
Koala lives in any part of the world. . . . 

Immense numbers of Koalas must have been destroyed by those young 
"sportsmen" of an earlier- generation than ours, but there seems never to have 
been any regular hunting with a view to marketing the skins. Yet the fur 
is very thick and warm, and, I am told, is in great demand by men living in 
Northern Canada and Europe .... 


Apart from the shooting which so greatly reduced their numbers, I firmly 
believe that the next most important factor was the bush fires which, during 
the last twenty or thirty years have ravaged practically the whole of this 
State. . . . The Koala falls an easy victim. . . . 

Between twenty and thirty years ago, some fishermen living at Corinella 
took a few Native Bears across to French Island, where . . . they thrived and 
multiplied. From this island they were introduced to Phillip Island where 
they are now one of the principal attractions to tourists. 

Despite the drawback of practically annual fires in the scrub, 
the Koalas "were holding their own on French Island until rabbits 
were introduced." Cats were then liberated to cope with the latter, 
but attacked the bird life, and consequently insect pests multiplied 

"The residents, noticing the trees dying, blamed the Koalas," 
quite without justification. 

"It became necessary then, in order to preserve the Koala, to 
select some other place for it, and the Fisheries and Game Depart- 
ment chose Quail Island, a Government reserve and sanctuary . . . 
in ... Western Port Bay. To this retreat some two or three 
hundred Koalas have now been transferred. ... It is hoped that 
on the three islands in Western Port the Koalas will have a safe 
home." (Lewis, 1934, p. 75.) 

Kershaw (1934, pp. 76-77) writes as follows concerning the sanc- 
tuary on Wilson's Promontory in southern Victoria: 

Totally unsettled, densely timbered, and, until recent years, rarely visited 
except by cattle musterers, this area has always been an ideal sanctuary. 
Thirty years ago the Koala was fairly numerous in spite of the periodical 
raids of skin-hunters. . . . 

Following the permanent reservation of the Promontory in 1908 as a National 
Park and Sanctuary for the preservation of the native fauna and flora, these 
interesting animals were no longer molested .... As a result Koalas gradually 
increased in numbers .... 

Their immunity from interference of any kind . . . resulted in their mul- 
tiplying to such an extent as seriously to threaten the existence of their 
natural food plant [Manna Gum, Eucalyptus viminalis]. . . . Quite a number 
of the trees had died. . . . 

Action was at once taken to reduce their numbers. Where it was possible, 
many were transferred to other parts of the Park, but in remote localities, 
such as Oberon Bay, transport was out of the question so that it became 
necessary, in order to save the remaining trees, to have a number destroyed. 
[Yet] in certain localities, this particular Eucalypt was practically exterminated. 

Their food-plant gone, many of the animals died, others worked back into 
the more heavily timbered ranges of the interior where they found suitable 
food among the Blue Gums. With a view to their acclimatization in some of 
the other States several Koalas were forwarded to New South Wales, South 
Australia, and Western Australia. 

Native Bears are still fairly numerous in the timbered country on the 
northern and eastern coasts of the Promontory and among the big timber 
in the vicinity of Sealer's Cove. 


The slight information available concerning the species in South 
Australia is summarized by Jones (1924, p. 187) : 

At one time the Koala was without doubt an inhabitant of South Australia, 
and many men now living can remember the time when it was by no means 
uncommon in certain districts of the South-eastern portion of the State. No 
more than ten years ago Koalas have been killed well within the geographical 
limits of South Australia. If it inhabits South Australia to-day is rather 
doubtful, although reliable information would point to the fact that a 
remnant of the stock may still linger not far from the Victorian border. So 
far as I know no example of the South Australian race has been examined 
scientifically, and no specimens seem to have been preserved. Victorian 
animals were liberated on Flinders Chase, Kangaroo Island, in November, 
1923, and it is hoped that they will become established in that faunal 

The Koala is completely protected by law in Victoria. 

Family VOMB ATID AE : Wombats 

The two currently recognized genera of wombats, represented by 
six forms, are confined to eastern and southern Australia, Tasmania, 
and islands of Bass Strait. Four subspecies are treated here. 

Island Wombat; Flinders Island Wombat 


Didelphis Ursina Shaw, Gen. Zool., vol. 1, pt. 2, p. 504, 1800. (Presumed by 
Spencer and Kershaw (19106, p. 39) to be based upon the "Wombach" of 
Hunter, in Bewick, Hist. Quadrupeds, ed. 4, p. 522, 1800. Type locality 
"New Holland" = Clarke Island, Bass Strait, according to Spencer and 
Kershaw (19106, pp. 37-39) ; but Cape Barren Island, Bass Strait, accord- 
ing to Iredale and Troughton (1934, p. 34).) 

FIGS.: Peron and Freycinet, Voyage Terres Australes, atlas, ed. 1, pi. 28, 
1811, and ed. 2, pi. 58, 1824; Cabrera, 1919, pi. 17, fig. 1. 

Formerly an inhabitant of several of the larger islands of Bass 
Strait, this Wombat has been exterminated on all of them except 
perhaps Flinders Island. It is also represented by a small colony 
introduced at Eddy stone Point, Tasmania. 

This is the smallest of the Wombats; hair coarse, varying from 
light sandy brown to blackish; rhinarium naked. Head and body, 
775 mm. Weight, 25-30 pounds (Spencer and Kershaw, 1910a, p. 29) . 

This species was originally known from King, Deal, Cape Barren, 
Clarke, and Flinders Islands in Bass Strait. At the time of its 
discovery, about 1798, its numbers were evidently considerable. 
Flinders (1814, vol. 1, p. cxxxv) found it more numerous on Cape 
Barren Island than on Clarke Island; he reports it as "commonly 
seen foraging amongst the sea refuse on the shore." 

Home (1808, p. 304) gives an entertaining description of an 


individual secured on Flinders' voyage and kept for two years as a 
pet in a house in England. It appeared intelligent as well as at- 
tached to its human friends. 

In their account of King Island, Peron and Freycinet (1816, 
vol. 2, p. 14) describe the local Wombat as a gentle and stupid 
animal, valuable for its delicate flesh. They also give an interesting 
picture of its tractability. They say it had been reduced to a domes- 
tic state by some English fishermen, going by day into the forests 
to seek its food, and returning in the evening to the cabin which 
served as its retreat. 

Spencer and Kershaw (19106, p. 48) write as follows: 

It is many years ago since the King Island wombat was exterminated. 
When the island was visited by a party of the Victorian Field Naturalists 
Club in 1887, no trace of it was discovered nor, during the process of clearing 
the land that has been vigorously carried on during recent years, has any 
record of a living wombat been made. 

Flinders Island afforded the only prospect of securing a living specimen 
of the Bass Strait species. [In 1908] a considerable part of the north, north- 
east, and north-west coast line was examined, and abundant evidence was 
obtained to prove that the animal, though very rare and difficult to obtain, 
was not extinct. In the deserted hut of a half-caste native at Killiecrankie 
two skins were found. ... On the island there are, in addition to a few 
settlers, a number of half-castes .... The existence of the wombat is well- 
known to them, but it is by no means easy to secure. ... On Cape Barren 
Island . . . the animal was found to be quite extinct, though well-known 
under the name of "badger" .... 

The animal is now extinct everywhere except on Flinders Island. 

An animal as large as a Wombat, always limited in numbers by 
an island habitat, could scarcely be expected to survive indefinitely, 
when confronted by deforestation as well as by the presence of 
settlers and half-castes who evidently prized its flesh. 

It is "now believed to be represented by small colonies on Flinders 
Island. Observation and careful provision for their safety may be 
necessary to avoid extinction." (E. Le G. Troughton, in litt., April 
16, 1937.) 

"The Flinders Island wombat has been introduced, and there is a 
small colony ... at Eddystone Point, North-East Tasmania. They 
were liberated there by the lighthouse-keepers." (Lord, 1928, p. 20.) 

[The Tasmanian subspecies, Vombatus ursinus tasmaniensis 
(Spencer and Kershaw), "has always and still does exist in large 
numbers in Tasmania" (R. Boswell, in litt., May 13, 1937).] 

[The common Wombat (Vombatus hirsutus hirsutus (Perry)) is 
still more or less numerous in wild and rugged portions of south- 
eastern Queensland, New South Wales, and Victoria. In southeastern 
South Australia another subspecies, Vombatus hirsutus niger 
(Gould), has been recognized; but no information concerning its 
numerical status is at hand.] 



Hairy-nosed Wombat 


Phascolomys latifrons Owen, Proc. Zool. Soc. London 1845, p. 82, 1845. ("Con- 
tinental (South) Australia.") 

FIGS.: Gould, 1863, vol. 1, pis. 59, 60; Wolf, 1867, vol. 2, pi. 27; Royal Nat. 
Hist., vol. 3, p. 266, fig., 1894-95; Jones, 1924, p. 267, fig. 189. 

This Wombat is now practically restricted to coastal South 
Australia, though once extending a little farther east and west; its 
numbers have evidently been severely reduced. 

FIG. 9. Hairy-nosed Wombat (LasiorhiniLs latifrons latifrons). 
After Wolf, 1867. 

The fur is soft and silky; general color grizzled gray, somewhat 
dappled; chin dark; cheeks, throat, and chest white; belly gray; 
ears long and narrow; rhinarium hairy; tail rudimentary. Head 
and body, 900 mm. (Jones, 1924, pp. 266-267.) 

In South Australia the species has been recorded from Mount 
Gambier, Port Augusta, Port Lincoln, River Murray, River Light, 
Fowler's Bay, Yorke Peninsula, Blanchetown, Blyth, 30 miles 
north of Adelaide, and Nullarbor Plain. "Apparently its distri- 
bution does not extend into the more northern parts of South 
Australia" (Spencer, 1896, p. 3). Specimens are recorded from 
Eucla, in the extreme southeast of Western Australia (Jones, 
1924, p. 268). E. Le G. Troughton writes (in litt., April 16, 
1937) that it was once plentiful, according to early observers, 
in southwestern New South Wales and Victoria, but now is ap- 
parently restricted to coastal South Australia, the inference 
being that survival is not assured. A number of specimens were 


recorded by Kershaw (1909, p. 118) from Deniliquin, N. S. W., 
close to the Victorian border. 

"Although in some parts of the colony [South Australia], es- 
pecially on Yorke's Peninsula and about Port Lincoln, the holes of 
these Wombats are very numerous, yet the animals are but rarely 
seen. Many of the oldest colonists have informed me that they 
never saw a Wombat alive. . . . The flesh they [the blacks] de- 
scribe as being like pork, and excellent eating." (Gould, 1863, vol. 1, 
p. 68.) 

"It could probably hold its own under present day conditions, and 
with existing introduced enemies, if only it had adequate protection 
from man. That Wombats are harmless to small holders is not con- 
tended. So bulky an animal which drives tunnels with such ease 
is not, of course, desirable in closely settled or intensively worked 
agricultural areas. But South Australia possesses vast tracts where 
Wombats might burrow and live without detriment to any human 
enterprise. In these areas they need protection from man alone." 
(Jones, 1924, p. 270.) 

"Being in grave danger of extermination, and having a distribu- 
tion restricted to South Australia, it is the intention of the [Fauna 
and Flora] Board to attempt to acclimatise the wombat on Kangaroo 
Island; the sending of a single specimen to the Chase on October 1, 
1926, may therefore be recorded; others will be forwarded as soon 
as obtained" (Waite and Jones, 1927, pp. 323-324). 

On the other hand, H. H. Finlayson (in Hit., March 20, 1937) 
considers the species still "plentiful in a restricted habitat." 

Le Souef and Burrell (1926, pp. 293, 295) write as follows: 

The . . . hairy-nosed wombat is found in the drier inland areas; it also 
lives along the coast of the Great Australian Bight .... 

The hairy-nosed wombat has been killed out over a large part of its range. 
In the Riverina, where at one time it was fairly plentiful, the settlers had 
to get rid of it as part of the campaign against the rabbits, which pests had 
a very secure harbour in wombat burrows. 

The skin is not put to any commercial use, though the aborigines use the 
fur of Ph. latijrons for making string, coils of which are wound round their 

On its economic status E. Le G. Troughton remarks (in litt., 
April 16, 1937) : "Colonies were exterminated near settlement be- 
cause of damage to fencing and crops, and risk of injury to stock 
in the burrows." 

Southern Queensland Hairy-nosed Wombat 


Phascolomys gillespiei De Vis, Annals Queensland Mus., no. 5, p. 14, pis. 9, 10, 

1900. (Moonie River, southeastern Queensland.) 
FIG.: De Vis, 1900, pi. 10. 

1 For the use of this combination, see Longman, 1939, p. 286. 


This Wombat is "apparently now extremely rare and restricted 
to remote parts of large properties." Its extinction is "apparently 
imminent." (E. Le G. Troughton, in litt., April 16, 1937.) 

General color gray, mixed with black, and washed with fawn 
(especially on rump and back) ; inner surface of ears, throat, and 
chest white; a broad curved blotch before and a spot behind the 
eye, black; forearm and feet dark brown; rhinarium hairy; skull 
relatively broader than in other Wombats. Head and body, 1,020 
mm. (De Vis, 1900.) 

The existence of a Wombat in Queensland was regarded as more 
or less mythical until three specimens of the present form were 
secured at the type locality and vicinity in the last decade of the 
nineteenth century. The subsequent record of the animal seems 
very meager. 

In 1923 Wilkins (1928, pp. 25-27) made a search for it in the 
Moonie River district, near Hollymount, finding "ancient tunnel- 
lings of many wombats" but not encountering any of the animals. 
He concluded that "there is no doubt that it is almost, if not quite, 
extinct in this district." 

Its range would appear to be restricted to southeastern Queens- 

Central Queensland Hairy-nosed Wombat 


Lasiorhinus latifrons barnardi Longman, Mem. Queensland Mus., vol. 11, pt. 3, 
p. 283, 1939. ("Epping Forest Station, 75 miles west of Clermont," east- 
central Queensland.) 

This recently described Wombat is known from four specimens 
(three of which are only skulls) , and it is considered on "the verge 
of extinction" (Longman, 1939, p. 287) . 

General dorsal color brown, mottled with gray, and interspersed 
with black hairs; rhinarium completely clothed with short brown 
hairs; ears elongate, well haired outside, with white tufts; under 
parts dirty gray. Total length, 3 feet 4 inches; tail, 2^ inches. 
(Longman, 1939, pp. 283, 286.) 

Although Wombats "were widely distributed in Queensland in the 
Pleistocene and two present-day species were known to occur spar- 
ingly in southern parts of the State, it was somewhat surprising 
to have definite evidence of living wombats in a locality in central 
Queensland. This extends their range by over 400 miles. . . . 

"Mr. Charles Barnard reports that there were many burrows in 
the district, but very few tracks of the animals were seen. . . . 

"Only three animals were seen, one of which was shot. As sug- 


gested by Messrs. Barnard, it is probable that these wombats were 
much more numerous in earlier years, but successive periods of 
drought have brought them to the verge of extinction. . . . The 
specimen shot . . . has been feeding on ... stems and leaves, 
including awns of the Comet River Grass, Perotis rara" (Longman, 
1939, pp. 283, 286-287.) 

Longman adds a report of Wombats seen distinctly about 1917 
in the Tambo district, south-central Queensland. 

Family MACROPODIDAE: Kangaroos, Wallabies, etc. 

This largest of marsupial families contains approximately 19 
genera and 125 forms. It ranges through Australia, Tasmania, New 
Guinea, and neighboring islands. Accounts of 27 forms appear in 
the following pages. 

St. Francis Island Rat-kangaroo 


This extinct animal, a former inhabitant of one of the islands in 
the Great Australian Bight, does not seem to be represented in the 
museums by so much as a skeletal fragment upon which a technical 
name might be based. Its brief and tragic history is recounted by 
Jones (1924, pp. 214-215) : 

Upon St. Francis Island in Nuyts' Archipelago there lived, during the 
time of the present occupiers, large numbers of what was evidently a species 
of Bettongia. Since the mammalian fauna of the islands of the Bight has 
proved, in so many instances, to exhibit distinctions from the types inhabiting 
the mainland, it is worth while recording what can still be ascertained con- 
cerning this interesting and recently exterminated animal. 

When the island was first settled, some forty years ago, "Rat-Kangaroos," 
or "Tungoos" were swarming. The animals do not seem to have formed 
burrows, but they lived in the undergrowth, and used frequently to hop into 
the homestead to take bread or other eatables thrown to them from the 
table. They do not appear to have been nocturnal; they do not seem even 
to have been afraid of the human invaders of the island. Their only offence 
seems to have been that they had a liking for the garden produce of the 
family who settled on the island. 

Cats were introduced in order to exterminate the Tungoos, and their work 
has been done completely. To what species the animal belonged can never 
be known and the fact of its extermination in this manner is much to be 

There are many islands in the vicinity of St. Francis to which some 
members of the original colony could have been transported, and so given a 
chance to survive. 

The story is one of importance from the point of view of legislation for 
the protection of insular faunas, since it demonstrates clearly how rapidly 
and how completely an interesting island fauna may be destroyed and lost 
to science for ever. 


It is much to be hoped that Isoodon nauticus, Petrogale pearsoni, Thylogale 
flindersi, Leporillus jonesi, and Rattus murrayi are not permitted to follow 
the Tungoos of St. Francis Island into the ranks of recently exterminated 

Gaimard's Rat-kangaroo 


Kangurus Gaimardi Desmarest, Mammalogie, vol. 2, suppl., p. 542, 1822. 

(Vicinity of Port Jackson, New South Wales.) 
FIG.: Quoy and Gaimard, Voy. Uranie et Phys., Zool., atlas, pi. 10, 1824 

(as Hypsiprymnus white}. 

This rat-kangaroo of eastern Australia is apparently extinct. 

The general color is grizzled gray, with a yellowish tinge; tail 
colored like body for the basal third, then darkening and the hair 
lengthening until there is a distinct black crest on the terminal third ; 
under side of tail white. Head and body, 390 mm.; tail, 280 mm. 
(Thomas, 1888, p. 109.) 

Le Souef remarks (1923, p. 110) that this is one of three mammals 
that "are entirely confined to the fox area of Eastern Australia" and 
"require our immediate attention if the remnants are to be saved. 
... I cannot locate any Gaimard's Rat-Kangaroos; they used to 
live on the Mountains and western plains of New South Wales." 

Le Souef and Burrell say (1926, p. 233) : "We have noted Gaim- 
ard's rat-kangaroo in the open forest on the Blue Mountains . . . ." 
They add, in regard to the group of rat-kangaroos in general: "Be- 
fore the advent of the fox the rat-kangaroos were extremely numer- 
ous, so much so that special measures had to be taken by settlers 
to protect crops and haystacks, but now many species throughout a 
large part of their range are very rare, and presumably in a short 
time they will be a thing of the past wherever the fox can pene- 

"I think that this species is definitely extinct. I have not seen or 
heard of it for upwards of 20 years." (A. S. Le Souef, in Hit., 
February 15, 1937.) 

E. Le G. Troughton writes (in Hit., April 16, 1937) that it once 
inhabited coastal New South Wales, but it is now apparently extinct, 
possibly since the advent of the fox. 

According to C. W. Brazenor (in Hit., March 3, 1937) , there are 
few Victorian specimens, and the last record was in 1877. 

Longman (1930, p. 59) includes southern Queensland in the range 
of the species. 


Gray's Rat-kangaroo 


Hypsiprymnus Graii Gould, Proc. Zool. Soc. London 1840, p. 178, 1841. 

("Swan River," Western Australia.) 
FIGS.: Gould, 1855, vol. 2, pi. 64; Cabrera, 1919, pi. 14. 

This subspecies of the Western Australian mainland has suffered 
pronounced restriction of range and reduction in numbers. 

Fur long and soft; general color above (including back of ears) 
ashy brown; sides of head and body very faintly tinged with yel- 
lowish ; under parts dirty white ; feet very pale brown ; tail brown, 
except the terminal third, which is covered with longish white hairs. 
Head and body, 457 mm. ; tail, 292 mm. (Gould, 1841c, pp. 178-179.) 

Gray's Rat-kangaroo is apparently now confined to a compara- 
tively small area in the southwest of Western Australia. Short- 
ridge (1910, p. 823, map) indicates a former distribution covering 
almost the entire southern half of that state. The line of demarca- 
tion or intergradation between this subspecies and B. I. harveyi re- 
mains undetermined. 

Gould (1863, vol. 2, p. 74) "received examples of this animal from 
various parts of the south-western coasts of Australia, and it appears 
to be ... abundant in the plains ... in the neighbourhood of 
Perth in Western Australia." He quotes Gilbert to the effect that 
"it is one of the most destructive animals to the garden of the settler 
that occurs in Western Australia, almost every kind of vegetable 
being attacked by it, but especially peas and beans." 

Thirty-five years ago it was "very abundant in many parts of the 
South-West, differing curiously from the insular form in not occur- 
ring near the coast." It did not then appear "to exist on the main- 
land to the north of the Swan River." Specimens were recorded from 
Arthur River, Woyaline Wells, Boyadine-Dale River, and Dwala- 
dine. (Shortridge, 1910, pp. 822-823, fig. 258.) 

More recently "this species, which was once very common in the 
interior, is now confined to the Great Southern area between Beverley 
in the north and Kojonup in the South" (Glauert, 1933, p. 26). 

The reduction in range suggests little hope for survival unless 
there exists a suitable reserve from which foxes can be excluded 
(E. Le G. Troughton, in Hit., April 16, 1937) . 

[On the islands of Sharks Bay, Western Australia, occurs the 
typical subspecies, Lesueur's Rat-kangaroo (B. I. lesueur (Quoy and 
Gaimard)), which has survived in considerable numbers, probably 
owing to the protection afforded by an insular habitat. Glauert 
(1933, p. 26) extends the range of this form far north along the 
coast of Western Australia: "Years ago the animal was common 


near Roebuck Bay (Broome), where K. Dahl obtained numerous 

Harvey's Rat-kangaroo 


Perameles Harveyi Waterhouse, Proc. Zool. Soc. London 1842, p. 47, 1842. 

("Port Adelaide, South Australia.") 
FIG.: Jones, 1924, p. 207, fig. 153. 

"In certain districts it is still by no means rare, but its decrease 
in numbers has been so rapid during the past twenty years that 
probably the remnant still existing must not be regarded as a very 
long lived one" (Jones, 1924, p. 207) . 

Fur dense and soft; general color above brown, penciled with 
white; sides of head and body tinged with yellowish; under parts 
impure white; tail rich brown above, dirty white below, the hairs 
becoming longer and white toward the tip (Waterhouse, 1842, p. 47) . 
The animal is shaped like a little stoutly built kangaroo and is about 
the size of a rabbit, with a short blunt head and little round ears. 
Head and body, 370 mm.; tail, 300 mm. (Jones, 1924, pp. 207, 210.) 

The exact distribution of this subspecies does not seem to have 
been determined. It originally ranged widely through South Aus- 
tralia, having been recorded from Adelaide, Port Lincoln, Gawler 
Plains, Lake Phillipson, the vicinity of McDoualFs Peak, and the 
northwest (Thomas, 1888, p. 113; Jones, 1924, pp. 210-211). It may 
be this form that extends also into Central Australia, but the area 
where it approaches or intergrades with B. L graii apparently re- 
mains to be ascertained. 

Gould (1863, vol. 2, p. 74) recorded it as "abundant in the plains 
around Adelaide." 

Spencer records "Bettongia lesueuri" from Central Australia, but 
he lists no specimens and gives it the native name of "Mal-la," 
whereas Finlayson (1935, p. 62) applies the native name "maala" 
to Lagorchestes hirsutus of the same general region. 

Spencer's account (1896, p. 16) is as follows: 

"This is the common sand-hill rat-kangaroo of Central Australia, 
and is perhaps . . . the most common form of marsupial amongst 
the sandy plains and sand-hills .... 

"We found it during the whole course of the [Horn] expedition, 
and there can be no doubt but that it is distributed right across 
South, Central and West Australia." 

As with so many other Australian species, we owe the chief 
account of this animal's status and life history to Jones (1924, pp. 
210-211) : 

This Rat Kangaroo, which is probably the only living representative of 
the Sub-family left in South Australia, is still existing in some numbers in 


certain districts in the North- West. Here it lives in company with the 
rabbits, sharing the larger warrens with them .... The choice of a warren 
seems largely to be determined by the quantity and nature of the herbage 
in the neighbourhood, for in these waterless districts Rat Kangaroos are 
dependent on the succulent sand hill vegetation. Rabbits are so universally 
spread over the country that there probably does not exist to-day a Bettongia 
colony living in its own burrows. It has thrown in its lot with the rabbit, 
and although it appears to have its own appartments [sic] in the complicated 
system of the large warrens, it is merely a tenant, forming a part of a 
community in a manner which is rather remarkable when its exceedingly 
pugnacious character is considered. Nevertheless, though it lives in apparent 
harmony with the rabbits, and avails itself of the shelter of their burrows, 
it is suffering for the partnership. The remnant of the Tungoos is living in an 
environment in which there is a severe competition for succulent food. In 
good seasons there is enough juicy herbage for cattle and rabbits as well as 
Rat Kangaroos but in bad seasons the rabbits and the marsupials perish 
in large numbers. Such losses among the rabbits are soon made good, but 
with the marsupials this is not the case, and probably the end of the Tungoo 
is not far off. When times are bad, and when the cattle and rabbits have 
eaten all the herbage of the sand hills, the Tungoos become extremely bold, 
and will enter a homestead in their search for anything to eat. They will 
come into a room and boldly face a cat in order to obtain some potato peelings ; 
they will scramble over a paling fence four or five feet high in order to get 
at the vegetable garden. They are bold and enterprising little animals which 
have made, and are making, a brave struggle against what seems an almost 
inevitable extermination. In the more cultivated districts of the South, 
where food is in plenty, the wholesale scattering of poisoned pollard has led 
to their complete extinction. The poison cart has done its deadly work on 
the slowly-breeding Tungoo, although the rapidly-breeding Rabbit has sur- 
vived the ordeal. In the North they are steadily being pressed out of exis- 
tence by the competition for food. 

When we remember that their numbers in rabbit warrens, even near to 
towns, was a source of constant annoyance to rabbiters less than twenty 
years ago, we can realise how destructive to the native herbivorous fauna the 
wholesale spreading of poisoned grain has proved to be. Nor must we forget 
that the remnant which still struggles on in the North is now exposed to the 
ravages of the fox. 

Concerning the introduction of this rat-kangaroo on Kangaroo 
Island, South Australia, Waite and Jones say (1927, p. 323) : 
"Specimens bred and reared in captivity in Adelaide were liberated 
within the observation enclosure on the reserve and seem to be 
doing well. If, when they are turned out into the larger world, they 
can avoid the goana (Varanus) they should prosper." 

H. H. Finlayson (in litt., March 20, 1937) regards the species as 
a whole as common in Western Australia, the Center, and north- 
western South Australia. 


Brush-tailed Rat-kangaroo 


Bettongia penicillata J. E. Gray, Mag. Nat. Hist. (Charlesworth), vol. 1, p. 584, 
1837. (No type locality was stated in the original description, but Thomas 
(1888, p. Ill) lists the type specimen from "New South Wales.") 

FIGS.: Gould, 1841, pi. 14; Waterhouse, 1846, vol. 1, pi. 7, fig. 1; Gould, 
1852, vol. 2, pi. 61; Lydekker, 1894, pi. 9 (ssp.?). 

This eastern Australian subspecies is either very rare or extinct. 
(The two other recognized subspecies likewise come within the scope 
of the present report.) 

The general color is ashy brown, penciled with white and brownish 
black ; cheeks and throat faintly tinged with yellowish ; under parts 
dirty white; hands and feet pale brown; tail brown above, pale 
brown below, the apical third with a black dorsal crest. Head and 
body, 343 mm.; tail, 285 mm. (Waterhouse, 1846, vol. 1, p. 213.) 

The former range extended from the Dawson Valley, Queensland, 
to Victoria, but apparently only on the inner side of the coastal 

Most of the information on this animal comes from Gould (1863, 
vol. 2, p. 71), who had opportunities of studying it while it was 
still abundant. "The eastern parts of Australia, particularly the 
districts on the interior side of the ranges of New South Wales, 
constitute the true habitat of the species .... I observed it to be 
very abundant on the Liverpool Plains, and on the banks of the 
river Namoi, from its source to its junction with the Gwydyr; but 
between the ranges and the coast I did not meet with it." He adds 
that the natives rarely pass without detecting its grassy nest on the 
ground, and almost invariably kill the sleeping inmates by dashing 
their tomahawks or heavy clubs at it. 

It is "apparently not now found in Eastern Australia" (A. S. Le 
Souef, in Hit., February 15, 1937) . "It is now very rare or extinct 
in New South Wales and Victoria" (E. Le G. Troughton, in Hit., 
April 16, 1937). 

C. W. Brazenor (in litt., March 3, 1937) considers the animal 
extinct in Victoria, where the last record dates from 1857. There 
are few Victorian examples in the National Museum of Melbourne. 

According to Finlayson (1931, p. 89), "Bettongia penicillata was 
taken by Lumholtz on Coomooboolaroo [in the Dawson Valley, 
Queensland], but has now apparently quite disappeared from there, 
and is not known elsewhere in the valley." 

While no specific information concerning the causes of the dis- 
appearance of this rat-kangaroo seems to have been offered, prob- 
ably the fox is largely responsible. 


Gould's Rat-kangaroo 


Bettongia Gouldii J. E. Gray, List Specimens Mammalia Brit. Mus., p. 94, 
1843 (nornen nudum). ("Head of Gulph St. Vincent," South Australia.) 

Bettongia Gouldii Waterhouse, Nat. Hist. Mammalia, vol. 1, p. 219, 1846. 
("South Australia.") 

"As far as can be ascertained at present, this animal seems to 
have disappeared from South Australia" (Jones, 1924, p. 212). As 
far as known, it was confined to this state. 

The type specimen, a very immature animal, is the only one of 
this subspecies that seems to have been described in detail. The fur 
is brownish, penciled with black and yellowish white; under parts 
white, more or less suffused with yellow; tail rusty brown at base, 
the terminal half black both above and below (Waterhouse, 1846, 
vol. 1, p. 219). Head and body, 390 mm.; tail, 310 mm. (Thomas, 
1888, p. 111). 

Jones (1924, pp. 212-214) furnishes practically all the available 
information on the former and present status of this rat-kangaroo: 

It is possible that it may prove to be still living somewhere in this State, 
and if there is any hope of such a survival it would seem that the South-East 
or the extreme North-East holds out the greatest promise. 

Not only does it appear to have died out completely over the greater portion 
of the State, but no specimen of the South Australian form seems to have 
been preserved in the zoological collections in Australia. At present, so far 
as this State is concerned, the race is represented only by some half-dozen 
skulls. . . . 

Only a few years ago this animal was extremely common over the greater 
part of South Australia. Twenty years ago the dealers in Adelaide did a 
great trade in selling them by the dozen at about ninepence a head for coursing 
on Sunday afternoons. It may surprise people who remember those days to 
know that there is not a preserved specimen, not even a skin of the animal, 
available for scientific study in South Australia to-day. In the same way 
it will one day surprise the rising generation when they realise that the few 
native animals they are now familiar with are gone for ever. . . . 

It is much to be hoped that, should some remnant of the South Australian 
race be found still living in the more bush-covered portions of the South 
or of the North-East, steps will be at once taken that it may be preserved 
and protected by every possible means. 

Ogilby's Rat-kangaroo 


Hypsiprymnus Ogilbyi Waterhouse, Naturalists' Library (Jardine), vol. 11, 
Marsupialia, p. 185, 1841. ("Western Australia in the neighbourhood of 
Swan River"; Thomas (1888, p. Ill) lists the type specimen from "York, 
W. A.") 

FIG.: Gould, 1852, vol. 2, pi. 62. 

Although very plentiful in the southwest of Western Australia a 
generation ago, this subspecies is now "reduced in numbers" (L. 


Glauert, in Hit., March 17, 1937) , and concern may well be felt as 
to the possibility of its following the other two subspecies on the 
road to extinction. 

It differs from B. p. penicillata in its generally darker coloration; 
in the rusty red of the base and sides of the tail; in the rufous 
coloring of the feet; in the terminal half of the tail being black 
both above and below ; and in the longer ears and more slender tarsi 
(Water-house, '1841, p. 186; Gould, 1863, vol. 2, p. 72). Head and 
body, 360 mm.; tail, 310 mm. (Thomas, 1888, p. 111). 

Gould (1863, vol. 2, p. 72) quotes Gilbert's notes from Western 
Australia: "This species appears to be equally abundant in all 
parts of the colony, but to evince a preference, perhaps, for the 
white-gum forests. . . . This animal is one of the favourite articles 
of food of the natives, who are very quick in detecting the nest, and 
generally capture the little inmate by throwing a spear through the 
nest and transfixing it to the ground, or by placing the foot upon 
and crushing it to death." 

Shortridge (1910, pp. 821-822, map) found it "very plentiful in 
the South -West, where, unlike Bettongia lesueuri, it occurs near the 
coast, extending as far north as the Moore River, becoming very 
rare at its northern limit. Formerly recorded from Sharks Bay, 
as so many of the other South-Western marsupials have been. 

"Although getting scarce in the more settled districts, both species 
of Bettongia are sufficiently numerous in many places to be rather 
destructive to crops, on which account they are often trapped and 
poisoned off in large numbers." 

Shortridge records specimens from King River, Dwaladine, Woy- 
aline Wells, Yallingup, and Burnside. From Perth southward, ac- 
cording to Glauert (1933, p. 26), it "is found in the coastal area as 
well as inland to the Great Southern and beyond." E. Le G. Trough- 
ton remarks (in litt., April 16, 1937) that "survival there may be 
significant of the influence of the fox not yet being fully asserted." 

Rufous Rat-kangaroo 


Bettongia rujescens J. E. Gray, Mag. Nat. Hist. (Charlesworth), vol. 1, p. 584, 
1837. (Type locality not stated in original description, but Thomas (1888, 
p. 104) lists the type specimen from "New South Wales.") 

FIGS.: Gould, 1841, pi. 13; Gould, 1855, vol. 2, pi. 65; Le Souef and Burrell, 
1926, fig. 51. 

Once common over much of eastern Australia, this species has 
largely or entirely disappeared from Victoria and New South Wales, 
but it remains common in the Dawson and Fitzroy Valleys, Queens- 



The fur is long and coarse; color above grizzled rufescent gray; 
an indistinct white stripe crossing the sides just in front of hips; 
under parts grayish white; ears rather long, black on outer surface; 
tail thickly haired, pale gray above, white below. Head and body, 
520 mm.; tail, 380 mm. (Thomas, 1888, pp. 103-104.) 

Gould writes (1863, vol. 2, p. 75) : "The south-eastern portion of 
the continent is its true habitat; and it is almost universally dis- 

FIG. 10. Rufous Rat-kangaroo (Aepyprymnus rufescens) 

persed over New South Wales, both on the sea and interior side of 
the mountain ranges. I found it very abundant on the stony sterile 
ridges bordering the grassy flats of the Upper Hunter, and in all 
similar situations. . . . From its invariably seeking shelter in the 
hollow logs" when startled from its nest, it "easily falls a prey to the 
natives, who hunt it for food." 

Of its status on the Comboyne Plateau, New South Wales, Chis- 
holm says (1925, p. 73) : "Not here now, but I am informed by an 
early settler that years ago they were a great pest to the farmers, 
and had to be persistently poisoned. This animal appears to be fast 
becoming extinct, probably largely due to the depredations of the 


fox, as living their daylight hours in a nest on the ground they 
become an easy prey for this animal." 

In Victoria, according to C. W. Brazenor (in Hit., March 3, 1937) , 
the Rufous Rat-kangaroo was "once fairly common. Now rare but 
probably survives in more inaccessible parts of eastern Victoria. 
Last record 1905." It is completely protected by law in that state. 

Finlayson writes (1931, pp. 85-86) of its status in Queensland: 

This interesting animal, though highly characteristic of coastal Queensland, 
has received very little mention in recent years, and there has been no pub- 
lished data from which one might estimate its position in the fauna of that 

Strangely enough it was not taken by Lumholtz, though it must have oc- 
curred in many of the districts in which he worked .... It has been twice 
recorded from North Queensland by O. Thomas . . . , and by Lonnberg and 
Mjoberg from Carrington . . . , but without comment, and as each record was 
based on a single individual, it might be inferred to be comparatively rare. 

In the Dawson and Fitzroy Valleys, however, this is far from being the 
case, and it is widely spread over the whole area from sea level to the tops 
of the plateaus. It occurs in almost all types of country, both open and 
forested .... The banks of creeks and river flats are favourite resorts, and 
there are few such places which by systematic beating cannot be made to 
yield up a few. . . . 

Like most of the coastal species it has little resistance to drought, and will 
go to great lengths in excavating holes in dry creek beds to get down to water 
level. In January, 1929, the Cariboe Creek ceased to run at Thangool, and 
for miles the sandy bed thus exposed was criss-crossed with the pads of 
Aepyprymnus coming down at night to drink at pot holes of their own making. 

In the cattle country it is stated by squatters to have diminished considerably 
in recent years, and by them it is regarded with indifference. But round 
many of the newly-formed cotton settlements in The Callide Valley it is 
plentiful, and at Thangool and Biloela and other points on The Cariboe has 
become an unmitigated nuisance and is cordially detested by the struggling 
settlers. Its raids on the crops are determined and resourceful, and as no 
ordinary fence will bar them for long, poisoning is the only effective check. 
Scores of thousands have been killed in this way, and skeletons (few and 
far between in Museums) are littered thickly round the cotton plots. 

On the outlook for the preservation of the Rufous Rat-kangaroo, 
E. Le G. Troughton writes (in litt., April 16, 1937) : "There has 
been a marked shrinkage of the once abundant species in coastal 
N. S. Wales and Queensland, suggesting that this small, specialized, 
and rather open country species is unlikely to survive, except pos- 
sibly in northern coastal Queensland where it may favour less open 
country, and the fox may not become established." 

Gilbert's Rat-kangaroo 


Hypsiprymnus Gilbertii Gould, Mon. Macropodidae, pt. 1, text to pi. 15, 1841. 

("King George's Sound," Western Australia.) 
FIGS.: Gould, 1841, pi. 15; Gould, 1854, vol. 2, pi. 69. 



The annals of this species are brief and tragic. It was discovered 
in Western Australia in 1840 by John Gilbert and is represented by 
his two specimens in the British Museum, but it has never since 
been encountered in the flesh by a zoologist and is undoubtedly 

General color above mingled gray, brown, and black; central and 
lower part of back washed with reddish brown; a blackish median 
line from nose to forehead; under parts grayish white; tail black, 
thinly clothed with short hairs. Total length, 558 mm. ; tail, 158 mm. 
(Gould, 1841, pt. 1, text to pi. 15.) 

FIG. 11. Gilbert's Rat-kangaroo (Potorous gilbertii) 

Gould (1863, vol. 2, p. 79) quotes Gilbert's field notes as follows: 

This little animal may be said to be the constant companion of Halmaturus 
brachyurus, as they are always found together amidst the dense thickets and 
rank vegetation bordering swamps and running streams. The natives capture 
it by breaking down a long, narrow passage in the thicket, in which a number 
of them remain stationed, while others, particularly old men and women, 
walk through the thicket, and by beating the bushes and making a yelling 
noise, drive the affrighted animals before them into the cleared space, where 
they are immediately speared by those on the watch: in this way a tribe of 
natives will often kill an immense number of both species in a few hours. I 
have not heard of the Hypsiprymnus Gilberti being found in any other part 
of the colony than King George's Sound. 

Shortridge (1910, pp. 824-826, map) gives the following account: 

"It is quite possible that they [P. gilbertii and P. platyops] are 

now entirely extinct, although I picked up six old skulls of Potorous 


gilberti near the entrances of some caves in the Margaret River 
district, and they may still exist sparingly in that and other locali- 
ties, as they are very liable to be overlooked on account of their 
great external resemblance to Macropus brachyurus. 

"The animal known to natives as 'Wurrark' around the Margaret 
River is probably Potorous gilberti, said to frequent marshy country, 
and although formerly numerous, it is thought to have almost, if 
not entirely, died out. A few may still occur towards Cape Leeuwin." 

L. Glauert (in litt., March 17, 1937) considers the species extinct. 

Broad-faced Rat-kangaroo 


Hypsiprrymnus platyops Gould, Proc. Zool. Soc. London 1844, p. 103, 1844. 
("Swan River," Western Australia; the type, according to Waterhouse 
(1846, vol. 1, p. 232), is labeled as coming from "Walyema Swamps, about 
forty miles north-east of Northam, Western Australia.") 

FIGS.: Gould, 1851, vol. 2, pi. 70; Le Souef and Burrell, 1926, fig. 55. 

This Western Australian species, regarded as rare at the time of 
its discovery a century ago, and represented by apparently no 
more than three specimens all told, is evidently extinct. 

Face extremely broad and, with sides of body, brownish gray; 
back reddish brown; face and upper surface beset with numerous 
long yellowish-white hairs; under parts and limbs buffy gray; tail 
brown above, paler beneath. Total length, 482 mm.; tail, 177 mm. 
(Gould, 18446, p. 103.) 

"This species ... is so rare that an adult male in my own col- 
lection and another in that of the British Museum, both procured 
fin 1840] by Mr. Gilbert in Western Australia, one in the Walyema 
Swamps, near Northam in the interior, and the other at King George's 
Sound, are all the examples that have yet been seen" (Gould, 1863, 
vol. 2, p. 80) . 

"A single specimen from the Margaret River was sent to the 
London Zoological Society in 1908. This suggests that the species* 
still exists in that area" (Glauert, 1933, p. 26). 

Shortridge wrote in 1910 (p. 826) : "A small gregarious wallaby 
is said to have been at one time plentiful in the coastal scrub to 
the east of Albany; from the description it was probably one of 
these species [P. platyops and P. gikbertii]. It was known to the 
natives as 'Moort/ and according to them has entirely disappeared 
there. Described as being rather similar to Macropus brachyurus in 
habits, but more sluggish in its movements, on which account cats 
and bush-fires have probably caused its disappearance." 

A possible clue to the identity of the above-mentioned species 
appears in Gould's original description of platyops (18446, p. 103), 
wherein he cites the native name of "Mor-da," presumably current 



in the Walyema Swamps area. This bears a plausible similarity 
to the "Moort" of Shortridge, whereas the native name of gilbertii 
was "Grul-gyte" (Gould, 1841, text to pi. 15) or "Ngil-gyte" (Gould, 
1863, vol. 2, p. 79). 

For some years past the Broad-faced Rat-kangaroo has been 
considered possibly or probably extinct (Shortridge, 1910, p. 825; 
Le Souef and Burrell, 1926, p. 237; A. S. Le Souef, in litt., February 
15, 1937; E. Le G. Troughton, in litt., April 16, 1937). Finally, L. 
Glauert (in litt., March 17, 1937) definitely lists it as extinct. 

FIG. 12. Broad-faced Rat-kangaroo (Potorous platyops). After Gould, 1851. 

"Common" Rat-kangaroo; Long-nosed Rat-kangaroo; Dark 
Rat-kangaroo; Potoroo 


Didelphis tridactyla Kerr, Anim. Kingdom of Linnaeus, p. 198, 1792. (Based 

upon the "Kanguroo Rat" of Phillip, Voy. Botany Bay, p. 277, pi. 47, 

1789; type locality, "New South Wales.") 
FIGS.: Waterhouse, 1841, pi. 16, and Gould, 1854, vol. 2, pi. 67 (as Hypsiprym- 

nus murinus); Lydekker, 1894, pi. 8; Le Souef and Burrell, 1926, figs. 53, 

54; Finlayson, 1935a, pi. facing p. 99. 

Formerly ranging from South Australia through Victoria and 
New South Wales to southern Queensland, the Potoroo has become 
extinct in South Australia and possibly in New South Wales. Its 
status in Queensland does not seem to be very definitely known, 
but it survives in some numbers in certain districts of Victoria. 

The Potoroo is distinguished from other species of its genus by its 
elongated head and short tarsus; the fur is long, loose, and slightly 


glossy; general color dusky brown, penciled above with black and 
pale brownish yellow; naked part of rhinarium extending farther 
back than in P. platyops; under parts dirty yellowish white; tail 
clothed with short, stiff, black hairs, extreme tip white. Head and 
body, 393 mm.; tail, 235 mm. (Waterhouse, 1846, vol. 1, pp. 224- 

The brief history of the species in South Australia is discussed 
by Jones (1924, pp. 217-218) : 

The name "common" Rat-Kangaroo although that used in all books dealing 
with the marsupials, is a sadly inappropriate one. . . . 

Of the former distribution of this animal in South Australia no details can 
now be obtained. Save the bare record of its existence in this State [on the 
Murray River], which is given in the British Museum catalogue of 1888 
and which has been copied into all subsequent works, I know no other 
reference to the creature as a South Australian animal. . . . The remaining 
Potoroos should be carefully protected in those places where they still sur- 
vive, and efforts should be made for turning them down in properly safe- 
guarded sanctuaries. If this is not done there seems to be no doubt that 
the remnant of the stock will share the fate of the South Australian form and 
rapidly become extinct. 

Finlayson writes (19356, p. 221) concerning its status in Victoria: 

Few animals have been so obscure as to their status on the mainland as 
the Potoroo. Its former presence in the south-eastern district of this State 
[South Australia] is attested ... by the accounts of settlers, and by occa- 
sional bone fragments in cave deposits, but it does not seem to have been a 
common form west of the Glenelg [a river of southwestern Victoria], at the 
time of settlement. 

In Victoria, though better known than here, there have been few explicit 
references to it in the literature, which would enable one to judge as to how it 
was faring in the struggle for survival, until Mr. Brazenor, in 1933 stated that 
"though very uncommon it still persists ... in the north-eastern district, in the 
Grampians, and probably in the Otway Ranges," and he has since confirmed 
its presence in the last locality by personally collecting it there. 

I am able to add two other localities to these, viz., French Island in Western 
Port, and the Portland area in the western district, and to state that in the 
latter, at least, it is still plentiful. Its apparent scarcity is due, I believe, 
largely to its choice of dense undergrowth .... In 1927 a rabbit trapper, 
near Gorae, stated that he took over twenty of these "bandicoots" in a 
short season, and this I was able subsequently to prove, by overhauling the 
skulls at his dumps .... In the summer of the following year I took it 
myself near Heywood and had further reports of it, and again in the winter 
of the same year on French Island, and that no disaster has overtaken it 
since then is vouched for by several correspondents, and very recently (for 
the western district) by Professor Wood-Jones (in litt.}. 

C. W. Brazenor writes (in litt., March 3, 1937) that it was once 
common in eastern and southern Victoria but is now confined to 
small numbers in southwestern Victoria. He adds that it is com- 
pletely protected by law. 

Gould (1863, vol. 2, p. 77) gives an account of it under the name 
Hypsiprymnus murinus: "It is only in the swampy and damp parts 


of the brushes of New South Wales that the H, Murinus is to be 
found in any abundance. The district of Illawarra, Botany Bay, 
the low scrubs bordering the rivers Hunter, Manning, and Clarence, 
are the principal localities in which it may be successfully sought 

E. Le G. Troughton (in litt., April 16, 1937) refers to the main- 
land race as once common in the damp coastal regions of New 
South Wales, Victoria, and South Australia, but as now rare, no 
specimens having been received at the Australian Museum since 

Le Souef and Burrell remark (1926, p. 233) that the Potoroo, like 
all the members of the subfamily Potoroinae, makes for a hollow 
log when disturbed, and thus is often easily captured. 

Longman (1930, p. 59) records the species from southern Queens- 
land. Finlayson (1931, p. 89) did not find it in the Dawson Valley 
in that state. 

[The Tasmanian subspecies, P. t. apicalis (Gould) , remains com- 
mon in many localities (Lord, 1928, p. 19). Absence of the fox in 
Tasmania may render that country the only hope for the survival 
of any representative of the genus (E. Le G. Troughton, in litt., 
April 16, 1937).] 

Desert Rat-kangaroo; Plain Rat-kangaroo 


Bettongia campestris Gould, Proc. Zool. Soc. London 1843, p. 81, 1843. ("South 

FIGS.: Gould, 1851, vol. 2, pi. 66; Finlayson, 1932, pis. 7, 8, and 1935, pis. 

facing pp. 97. 98. 

"His [Finlayson's] rediscovery of the living Caloprymnus was a 
romance of modern zoology. The great John Gould had received 
three specimens from somewhere in South Australia in 1843. These 
three specimens in the British Museum remained unique. Calo- 
prymnus seemed to be as dead as the Dodo: and then Finlayson, 
with the assistance of Mr Reese of Appamunna, produced [in 1931], 
as a conjurer from his hat, living specimens of the long lost Plafti 
Rat-kangaroo." (Jones, in Finlayson, 1935a, p. 8.) 

Under fur dense and soft, very pale yellowish brown, the hairs 
tipped with sooty brown; interspersed with the under fur (especially 
on the back) are many long brownish white hairs, tipped with 
blackish; sides dirty yellowish; under parts dirty white; feet and 
tail very pale yellowish brown. Head and body, 400 mm.; tail, 
355 mm. (Gould, 1843, p. 81.) 

"Imagine a little animal about the bulk of a rabbit, but built 
like a kangaroo, with long spindly hind legs, tiny forelegs folded 


tight on its chest, and a tail half as long again as the body but not 
much thicker than a lead pencil, and you have it in the rough. But 
its head, short and blunt and wide, is very different from that of 
any kangaroo or wallaby, and its coat is uniformly coloured a clear 
pale yellowish ochre exactly like the great clay-pans and flood 
plains." (Finlayson, 1935a, p. 102.) Head and body, 254-282 mm. ; 
tail, 307-377 mm. (Finlayson, 1932, p. 165). 

The exact locality from which Gould's original specimens came 
(through Sir George Grey) is not now ascertainable; he merely 
stated (1863, vol. 2, p. 76) that "the stony and sandy plains of the 
interior of South Australia partially clothed with scrub are its 
native habitat." Only the recent range of the animal can be given 
with any precision. "Its proved distribution may be extended over 
a large area of the eastern portion of the Lake Eyre Basin, speci- 
mens and reliable records from observers personally known to the 
author having been obtained from as far south as Lake Harry and 
as far north as Coorabulka in South-West Queensland. The north 
and south limits of its range, as at present ascertained, are, there- 
fore, approximately, lats. 23 40' and 29 21' south. No records 
have so far been obtained west of Lake Eyre and the Kallakoopah, 
and the furthest easterly occurrence is at Innamincka, on the Barcoo, 
in long. 140 49' east." (Finlayson, 1932, p. 148; map, p. 149.) 

The recent history of the species may be summarized in Finlay- 
son's own words (1932, pp. 150-165) : 

In ordinary years the Lake Eyre Basin is a most unattractive area from 
the point of view of the mammal collector, and the disappearance of 
Caloprymnus from scientific ken must be attributed rather to lack of systematic 
collecting than to any sudden change in the status of the animal in the 
fauna, following Grey's discovery. All the evidence obtained by questioning 
blacks goes to show that in all probability it has had an uninterrupted tenure 
of the country, but it is equally certain that in normal times its numbers 
are small, since men like Mr. Reese, whose opportunities for observation are 
practically continuous throughout the year, affirm that in thirty-five years 
they have seen no more than twenty specimens. . . . 

At the time of my passage through the area, conditions as they bear upon 
animal life were very favourable and quite supernormal as compared with 
average conditions over a series of years. A period of seven years of drought 
had been broken, and vegetation had been restored on a comparatively 
lavish scale. All species of mammals were undergoing a quick increase in 
numbers, and rodents especially had assumed plague proportions. Most of the 
specimens of Caloprymnus were obtained, and the bulk of the observations 
upon it were made, on two flats lying east and west of Cooncherie Sandhill. . . . 

The numbers occurring in this particular locality were very considerable. 
In the course of a week's riding on the two flats over an area of perhaps 20 
square miles, 17 Oolacuntas were sighted. ... All the evidence obtained 
so far goes to show that its distribution at present is highly discontinuous 
but that it follows in a general way the fringes of the gibber plains .... 

The accounts of its feeding habits given by the blacks, and several other 


items of evidence, however, would point to its being largely phytophagous, 
or at least less rhizophagous than Bettongia, Potorous, and Aepyprymnus. . . . 
Where Diprotodon failed [to survive], Caloprymnus may yet succeed, but 
all the evidence of its physical structure is not more eloquent of changed 
conditions [from relatively humid to an arid climate] than its pathetic 
clinging to its flimsy grass nest, in a fiery land where a fossorial habit has 
become the main factor in survival. 

The first specimen flushed by Finlay son's party was run down 
with a relay of horses after a chase of 12 miles. Others were taken 
in the same way, while a native captured two by hand after stealing 
up to their grass nests. 

E. Le G. Troughton writes (in litt., April 16, 1937) : "The fact 
that the otherwise defenceless animal is peculiarly suited to ex- 
tremely barren and remote desert areas may ensure survival and 
prevent exploitation as a rarity, but the spread of the fox, seen 
personally near Marree in 1920, use as food by the blacks, and 
variable seasons may continue range shrinkage to extinction." 

"Common" Hare-wallaby; Brown Hare-wallaby 


Macropus Leporides Gould, Proc. Zool. Soc. London 1840, p. 93, 1841. ("In- 
terior of Australia"; according to Thomas (1888, p. 84), the cotypes 
are from the "interior of New South Wales.") 

FIGS.: Gould, 1841d, pi. 12; Qould, 1859, vol. 2, pi. 57; Royal Nat. Hist., 
vol. 3, p. 246, fig, 1894-95. 

This species is "apparently doomed to extinction" in its last 
stronghold in New South Wales (E. Le G. Troughton, in litt., April 
16, 1937). 

It resembles the Common Hare of Europe in size and in texture 
of fur; forelimbs very small; above variegated with black, brown, 
and yellow; pale yellow on sides and about eyes; belly grayish 
white; forelimbs black on upper part. Head and body, 495 mm.; 
tail, 330 mm. (Gould, 1841a, pp. 93-94.) 

The former range included the interior of New South Wales and 
Victoria, and the Murray River region of South Australia. 

Gould writes (1863, vol. 2, p. 67) : "I have but little doubt that 
this animal enjoys a wide range over the interior of New South 
Wales; it certainly inhabits the Liverpool Plains as well as those 
in the neighbourhood of the Namoi and the Gwydyr, from all of 
which localities I have received numerous examples." He adds: "I 
usually found it solitary, and sitting close in a well-formed seat 
under the shelter of a tuft of grass on the open plains." 

" According to Krefft, this species is common in the level country 
between the Murray and Darling rivers" (Lydekker, 1894, p. 54). 

E. Le G. Troughton (in litt., April 16, 1937) considers it "now 



very rare, and apparently doomed to extinction in New South 
Wales owing to denudation in over-stocked country, also populated 
by rabbits, and the fox." In 1938 (p. 407) he refers to it as "either 
extinct or nearly so." 

In Victoria, according to C. W. Brazenor (in Hit., March 3, 1937) , 
there were two records, the last in 1869. He considers the species 
extinct in that state. 

"It is tolerably abundant in all the plains of South Australia, 
particularly those situated between the Belts of the Murray and the 
mountain ranges" (Gould, 1841J, text to pi. 12). 

FIG. 13. Brown Hare-wallaby (Lagorchestes leporides). After Gould. 

Jones writes (1924, pp. 222-223) of its status in South Australia: 
"I know of no preserved specimens of this formerly common 
animal from which a description may be written of the actual form 
which inhabited this State. . . . 

"In the British Museum catalogue of 1888 five specimens in the 
collection are recorded as being from South Australia .... I have 
been unable to obtain any evidence of its present existence in the 
State, and in all probability it is completely exterminated." 

Rufous Hare-wallaby; Western Hare-wallaby; "Whistler"; 
"Spinifex Rat" 


Lagorchestes hirsutus Gould, Proc. Zool. Soc. London 1844, p. 32, 1844. 

("York District of Western Australia.") 
FIG.: Gould, 1849, vol. 2, pi. 58. 

This animal has long since disappeared from a large part of its 
former range in the west of Western Australia but survives in 


indefinite numbers in the east along the South Australian border 
and likewise in the northwestern part of the latter state. 

General color of fur, especially on hind quarters and under parts, 
rich sandy buff; head and back grizzled with grayish white; body 
beset, especially posteriorly, with numerous long, rich rufous hairs; 
space about eye reddish buff; ears large, grayish brown externally; 
feet yellowish buff. Total length, 698 mm.; tail, 266 mm. (Gould, 
1844a,p. 32.) 

Stirling and Zietz (1893, pp. 154-155) record four specimens from 
Western Australia ; they were taken during the Elder Expedition "in 
the Porcupine grass (Triodia irritans) country, south of the Barrow 
Range, before the exploring party entered the Victoria Desert. . . . 

"Mr. Streich informs us that this animal appears to be numerous 
in the northern parts of the Victoria Desert, where it often falls the 
prey to the Wedge-tailed Eagle." 

In an anthropological report on the same expedition, Helms says 
(1896, pp. 240, 255-256) : 

The Blyth Range, Barrow Range, and Victoria Desert tribes inhabit "spini- 
fex country," where subsistence is difficult to maintain, and but for the 
numerously occurring Largochestes [sic] hirsutus . . . and some other small 
marsupials, it would probably be impossible for them to live in such desolate 
districts. It can scarcely be wondered at that the majority of them appeared 
lean and starvation-stricken. . . . 

The Largochestes is almost, if not totally, absent here [in the vicinity of 
Victoria Spring] ; and some 150 to 200 miles to the north, the eagle-nests, 
which we had previously noticed daily, disappeared, which at once proved 
the decrease of these small marsupials. . . . 

Throughout the greater part of the interior, as far as the Expedition 
went, their [the blacks'] principal flesh-food is supplied by the small marsupials 
that harbor under the Zn'ocfoa-tussocks, and are commonly called "spinifex 
wallabies" (Largochestes hirsutus), occurring abundantly in many places in 
the triodia-regiou. . . . The blacks are very expert at killing the animals with 
the "turtimbo," or short throwing-stick .... To enable them better to get 
at this game they constantly burn large patches of the "spinifex" grass. 

Shortridge (1910, p. 819; map, p. 820) considers the "mainland 
form almost, if not entirely, extinct. Said possibly to still occur 
very sparingly on sand-plains to the east of Beverley and York 
where within quite recent times it was fairly plentiful. 

"A single specimen was recorded from Hastings, near Kojonup, 
in 1896, by the Perth Museum." 

Glauert states (1933, p. 27) : "It has long disappeared from the 
York district, where the first specimens were collected by Gilbert, 
but survives in the desert country near the South Australian border, 
and along the Canning Stock Route." 

Finlayson (1935, pp. 63-67) gained experience with the species 
in the northwest of South Australia. "The maala [L. hirsutus], 
though common in the more westerly spinifex tracts, is not often 
taken east of the [Aboriginal] Reserves." Much time was spent in 


1932 in an unsuccessful search for it. "A year later, ... on the 
south side of the Musgraves, we learned from the blacks that there 
was a small colony of maalas in a spinifex patch ten miles south 
of Koonapandi." 

Finlayson then gives an account of hunting in this spinifex patch, 
about 10 miles square, with some blacks. Their favorite method 
of firing the country was utilized. As the fire drove the maalas out 
of the tussocks, their chance of dodging the throwing-sticks was 
slender. Those that escaped the fire by remaining in their burrows 
were hunted out after the fire had passed. A satisfactory series of 
specimens was secured. 

The Rufous Hare-wallaby "requires observation and close pro- 
tection wherever possible" (E. Le G. Troughton, in litt., April 16, 

[It is represented by doubtfully distinct insular subspecies on 
Dorre and Bernier Islands in Sharks Bay, Western Australia L. h. 
dorreae Thomas and L. h. bernieri Thomas. Evidently both are 
protected by their insular environment, and at least the latter is 
reported as plentiful (Shortridge, 1910, p. 819).] 

Banded Hare- wallaby; Banded Wallaby 


Kangurus Fasciatus Peron and Lesueur, in Peron and Freycinet, Voyage Terres 

Australes, vol. 1, p. 114, atlas, ed. 1, pi. 27, 1807. (Bernier Island, Sharks 

Bay, Western Australia.) 
FIGS.: Peron and Freycinet, Voyage Terres Australes, atlas, ed. 1, pi. 27, 

1807, ed. 2, pi. 57, 1824; Gould, 1842, pi. 30; Waterhouse, 1846, vol. 1, pi. 4, 

fig. 2; Gould, 1849, vol. 2, pi. 56. 

Although Shortridge says (in Thomas, 1907, p. 772) that he had 
never seen any animal, not even rabbits, in such numbers as this 
species on Bernier Island, it now seems a melancholy necessity to 
include it among the vanishing mammals of Australia. It is no 
longer common on the islands in Sharks Bay where it once swarmed, 
and it occurs in only a few isolated localities on the mainland of 
Western Australia. (The mainland form, sometimes recognized as 
L. /. albipilis (Gould) , is doubtfully distinct, and will not be treated 
separately here.) 

The animal is about the size of the Common Hare of Europe; 
fur very long and soft, brown-gray, variegated with rusty, black, 
and white; space about eye bright rusty; numerous narrow dark 
cross bands on the back, most conspicuous posteriorly ; on the upper 
parts and sides are very long interspersed white hairs; under parts 
dirty white; hind feet with long, harsh, brownish-white hairs spring- 


ing from sides of two larger toes. Head and body, 444 mm.; tail, 
279-305 mm. (Waterhouse, 1846, vol. 1, pp. 87, 90.) 

At the time of its discovery by Peron in 1801, the species occurred 
in great numbers on the islands in Sharks Bay (Bernier, Dirk Har- 
tog's, and Dorre). A little more than a century later Shortridge 
(in Thomas, 1907, p. 772) found the animals swarming on Bernier 
Island. "It has been a particularly dry season, and they were very 
thin. Food was evidently insufficient for them all, and dead speci- 
mens were lying about in all directions. It would seem that they 
have no natural enemies on the island; and they breed to such an 
extent that the island will carry no more, and in times of drought 
a number have to die." He adds (1910, p. 818) : "It may be noted 
that sheep had been temporarily introduced there, while in the 
south of Dirk Hartog there is a large sheep station, and the wallabies 
are said to have entirely left that end of the island." 

Glauert (1933, p. 27) reports the species as "not common" on the 
islands in Sharks Bay. 

On the mainland of Western Australia Gilbert found it in densely 
thick scrubs, where "thie only possible means of obtaining it is by 
having a number of natives to clear the spot, and two or three with 
dogs and guns to watch for it. ... The natives are in the habit 
of burning these thickets at intervals of three years, and by this 
means destroy very great numbers." (Gould, 1863, vol. 2, p. 65.) 

Thomas (1888, p. 182) recorded specimens from Wongar Hills, 
York, and Perth. 

Shortridge (1910, p. 818; map, p. 817) found it "existing in a 
few isolated localities to the east of Pinjelly and Wagin, and accord- 
ing to natives the Pellinup and Salt River districts in the neighbour- 
hood of the Stirling Ranges. 

"Plentiful enough in the restricted areas in which they occur, 
frequenting thick prickly scrub." 

He also remarks (pp. 818-819) on the "most sudden and unac- 
countable" disappearance of this and a number of other mammals 
in the Western, South-Eastern, and Central districts of Western 
Australia; it "is said to have been first noticed about 1880." Short- 
ridge continues: 

The above areas are now, with a few exceptions, entirely devoid of indigenous 
mammals. This is said partly to account for the way in which the natives have 
been disappearing from the Western and Central districts of late years. . . . 

The entire disappearance of so many species, over such large tracts of country, 
is generally considered to be due to some epidemic or disease .... It may be 
noted, however, that they have died out chiefly in the drier parts of the country, 
where, except for the introduction of sheep, there has been very little altera- 
tion in the natural conditions. Rabbits, although already very numerous in 
the Centre and South-East, have not yet found their way to the North- West. 

The mammals of the South- West, to about as far north as the Moore River, 
... are rapidly retreating before civilisation. . . . The burning of forests 


and general clearing of the country, together with constant raids of dogs 
and domestic cats, are among the chief causes. 

Glauert (1933, p. 27) records the mainland form as rare, occurring 
"in a few isolated localities to the east of the Great Southern 

E. Le G. Troughton (in Hit., April 16, 1937) regards the extinction 
of the mainland form as probable and states that suitable reserva- 
tions must be made if the fauna is to survive on islands large enough 
for commercial use. 

Bridled Nail-tailed Wallaby 


Macropus jraenatus Gould, Proc. Zool. Soc. London 1840, p. 92, 1841. ("In- 
terior of New South Wales.") 

FIGS.: Gould, 1841d, pi. 3; Waterhouse, 1846, vol. 1, pi. 4, fig. 1; Gould, 1849, 
vol. 2, pi. 54; Le Souef, 1923, pi. 15; Australian Zoologist, vol. 3, pt. 4, 
pi. 20, 1923; Le Souef and Burrell, 1926, fig. 48. 

This beautiful wallaby, perhaps never very common, is threatened 
with extinction in its ancient home in eastern Australia. 

Form slender ; fur soft and short ; general color gray ; a white cheek 
stripe ; sides of neck washed with cream ; a white stripe on each side 
extending from occiput over the shoulder to join the white of the 
under parts behind the arm insertion; space between these stripes 
blackish brown ; tail black above and below toward tip, with a small 
terminal nail; limbs whitish, darker on hands and feet. Total length, 
825 (female) to 1,104 mm. (male) ; tail, 380 (female) to 482 (male). 
(Gould, 1841a, p. 92, and 1841d, text to pi. 3.) 

Its former range extended from southern Queensland to Victoria. 

"0. fraenata inhabits the brigaloe-scrubs of the interior of New 
South Wales and Queensland, and probably South Australia" (Gould, 
1863, vol. 1, p. xxi). Gould (1863, vol. 2, p. 62) writes further of 
its occurrence: 

It is a native of the south-eastern portions of Australia, and the locality 
nearest to the colony of New South Wales in which I observed it was Brezi, 
on the river Mokai, whence it extended into the interior as far as I had an 
opportunity of proceeding; Mr. Gilbert subsequently discovered that it was 
common in the thick patches of scrub which are dispersed over all parts 
of the Darling Downs. It inhabits all the low mountain ranges, the eleva- 
tion of which varies from one to six hundred feet, and which are of a sterile 
character hot, dry, stony, and thickly covered with shrub-like stunted 
trees. . . . 

In the neighbourhood of Brezi the natives hunt this species with dogs, 
and often kill it with spears, bommerengs and other weapons; at Gundermein 
on the Lower Namoi I found myself among a tribe of natives who succeed 
in capturing them with nets .... 

Its flesh, like that of the other small Kangaroos, is excellent, and when 
procurable was eaten by me in preference to other meat. 



"There are probably only three species of animals that are entirely 
confined to the fox area of Eastern Australia. These require our 
immediate attention if the remnants are to be saved. They are the 
Bridle Nail-tailed Kangaroo . . . , the Brush-tailed Rock-Wallaby 
. . . , and Gaimard's Rat-Kangaroo. . . . 

"The only Bridle Nail-tailed Kangaroos that exist as far as 1 
know, are a few on Mr. Charles Baldwin's farm, near Manila, and 
some in Taronga Park. Attempts to get this species to live in a wild 
state in Taronga Park have failed, as they apparently cannot live 
in the tick area, their proper home being the foot hills of the Dividing 
Range of Eastern Australia." (Le Souef, 1923, p. 110.) 

FIG. 14. Bridled Nail-tailed Wallaby (Onychogalea fraenata). After Gould 
and photo by Berridge. 

"Occasionally one . . . will be picked up by the great wedge- 
tailed eagle. Remains of the animal have been found in the bird's 
aerie. . . . 

"This species, like so many other animals found in the more closely 
settled parts of Eastern Australia, is now becoming very scarce, and 
will probably soon be extinct. With the occupation of the land by 
sheep and cattle, and the competition of the rabbit, the food and 
shelter to which the wallabies were accustomed are decreasing. At 
the same time, their enemies are increasing as their possible living- 
grounds are becoming more and more restricted. But it is the 
imported fox that is making the wholesale clearance, threatening 
early extinction." (Le Souef and Burrell, 1926, pp. 210-211.) 

"It is not uncommon in some parts of Southern Queensland, and 
its pelts were frequently seen in the sales two or three years ago 


under the name of 'padmelon.' It is now a protected species." (Long- 
man, 1930, p. 59.) 

Finlayson reports (1931, p. 85) on its status in the Dawson Valley, 
Queensland: "Observed twice only, and no specimens obtained. It 
was obtained by Lumholtz in the Rockhampton district in 1880- 
1884, and recently Longman has stated that it is not uncommon in 
South Queensland. Over the greater part of the Dawson country, 
however, it is either absent or rare, as few reliable accounts of it 
could be obtained." 

"This gentle and beautiful species was once plentiful in inland 
N. S. W. south to the Murray River, and in coastal parts as far as 
Rockhampton in Queensland, but is now quite rare, or absent, over 
entire range. A colony has been established on a small river island, 
and such sanctuaries, free from foxes, probably represent the only 
means of preventing extermination." (E. Le G. Troughton, in Hit., 
April 16, 1937.) 

In Victoria there are a few records only, the last in 1867. The 
animal is now extinct in that state. (C. W. Brazenor, in Hit., March 
3, 1937.) 

"As an illustration of the rapid breeding of marsupials, the ex- 
perience of Mr. Chas. Baldwin, of Durham Court, Manilla, New 
South Wales, is illuminating. In eighteen months Mr. Baldwin, 
from five adults, bred seventy young of the bridled wallaby." (Hoy, 
1923, p. 166.) 

Crescent Nail-tailed Wallaby 


Macropus lunatus Gould, Proc. Zool. Soc. London 1840, p. 93, 1841. ("West 
coast of Australia"; Thomas (1888, p. 78) lists the type specimen from 
"Swan R., W. A.") 

FIGS.: Gould, 1849, vol. 2, pi. 55; Lucas and Le Souef, 1909, p. 78, fig. 

This wallaby is on the verge of extinction in the settled districts 
of Western Australia but survives farther east toward the Great 
Victoria Desert. 

In general appearance it is very similar to the Bridled Nail-tailed 
Wallaby but is slightly smaller; general color dark gray; face gray, 
a mark over eye and cheek stripe slightly paler; a prominent white 
crescent-shaped shoulder stripe, not encroaching on the neck; back 
and sides of neck rich rufous ; a whitish hip stripe and another stripe 
just above it; under parts whitish; tail uniform gray, its terminal 
nail as in 0. fraenata. Male: head and body, 500 mm. ; tail, 332 mm. 
(Thomas, 1888, pp. 77-78.) 

The former range of this species included the southern parts of 
Western Australia and Central and South Australia. 



"Mr. Gilbert's notes inform me that 'the Waurong ... is found 
in the gum forests of the interior of Western Australia, where there 
are patches of thick scrub and dense thickets . . . ; the dogs some- 
times succeed in driving it out to the open spots, when, like the 
Kangaroo rats, it runs to the nearest hollow log, and is then easily 
captured' " (Gould, 1863, vol. 2, p. 64). 


FIG. 15. Crescent Nail-tailed Wallaby (Onychogalea lunata) 

Shortridge (in Thomas, 1907, p. 768) considers it "very numerous 
in some localities" of Western Australia. He adds (1910, pp. 815- 
816, map) the following information: 

"Within a more limited area this species seems to have much the 
same range as Macropus eugenii, both forms frequently occurring 
together . . . not extending far, if at all, beyond Beverley in the 
North, or near the coast; its western boundary apparently being 
the Darling Range. 

"Also occurring in the southern interior of South Australia, where, 
however, it is little known and probably rare." 

Shortridge records 23 specimens from Arthur River and Woyaline 


"Mr. J. T. Tunney, whose fame as a collector is world-wide, in- 
formed me that I could only hope to get the Crescent Wallaby . . . 
along one obscure river [of Western Australia], and a forlorn hope 
at that. Such a Wallaby should be energetically sought, trapped, 
and placed, not in Zoological Gardens, but in the haven of a properly 
supervised national reserve." (Troughton, 1923, p. 155.) 

It occurs in "South-Western Australia, in isolated localities to the 
west of the lower Great Southern Railway, probably on the verge of 
extinction in the settled districts, but surviving further east towards 
'the Great Victoria Desert" (Glauert, 1933, p. 29) . 

It is "still found on the Nullarbor Plain" (A. S. Le Souef, in litt., 
February 15, 1937). 

Jones (1924, p. 234) writes of its status as follows: 

"In 1884 Mr. E. B. Sanger reported the Crescent-marked Wallaby 
from the Centre and in the British Museum catalogue of 1888 three 
South Australian specimens, collected by Sir George Grey, are 

"The Elder Expedition in 1891 met with it in the Everard Ranges 
.... The Horn Expedition of 1894 obtained two specimens at 
Alice Springs. I know of no more recent observations, and proba- 
bly so far as South Australia is concerned the animal has ceased 
to exist." 

C. W. Brazenor writes (in litt., March 3, 1937) of a single Vic- 
torian record, from the River Murray in 1857. 

"Regarded as verging upon extinction in the settled areas of its 
south Western Australian habitat, but surviving in the more desert- 
like conditions between the Great Victoria Desert and Trans-Rail- 
way, to the eastward. Extinction may be regarded as inevitable, 
without establishment under favourable conditions." (E. Le G. 
Troughton, in litt., April 16, 1937.) 

Doubtless settlement and the concomitant imported pests have 
accounted for the decline of this lovely wallaby. 

Brush-tailed Rock-wallaby 


Tufted-tailed or Mountain Kanguroo, K. pencillatiLs [J. E. Gray, in] Griffith, 

Smith, and Pidgeon, Anim. Kingdom (Cuvier), vol. 3, Mammalia, pi. 

opposite p. 49, 1827. (No type locality given.) 
Kangwrus Pencillatus * [J. E. Gray, in] Griffith, Anim. Kingdom (Cuvier), 

vol. 5, Mammalia, p. 204, 1827. ("New Holland" = "Sydney, N.S.W.," 

according to Iredale and Troughton, 1934, p. 42.) 
FIGS.: Waterhouse, 1841, pi. 22, and 1846, vol. 1, pi. 1, fig. 1; Gould, 1842, 

pi. 23, and 1853, pis. 39, 40; Lydekker, 1894, pi. 6; Le Souef and Burrell, 

1926, fig. 47. 

Formerly abundant in eastern Australia, this animal has suffered 
serious reduction of range and numbers. 

i Corrected to penicillatus in index volume, p. 23, 1835. 


The fur is long and thick; general color above dull brown, more 
rufous on rump; an indistinct black mark behind the shoulder suc- 
ceeded by a pale gray one; chin and chest pale gray; belly brown, 
tinged with yellow; anal region yellowish rufous; arms and legs 
rufous brown, becoming black at extremities; tail more or less 
bushy, basal part rufous, remainder black, tip sometimes yellow. 
Head and body, 720 mm.; tail, 560 mm. (Thomas, 1888, p. 67.) 

The Brush-tailed Rock-wallaby once inhabited the ranges of 
eastern Australia from southeastern Queensland to Victoria. 

In New South Wales, according to Gould (1842, text to pi. 23), 
"the species abounds wherever the kind of country suitable to its 
habits occurs. The specimens in my own collection were procured 
on the Liverpool range, and on the rocky sides of the mountains 
facing Yarrundi on the Dartbrook, a tributary of the Hunter. I also 
ascertained that it is very abundant on Turi, and the other moun- 
tains situated to the eastward of the Liverpool Plains, and it doubt- 
less ranges over a much greater extent of country than we are yet 
acquainted with. It is ... strictly gregarious, assembling in such 
numbers as to form well-beaten paths along the sides of the moun- 
tains they inhabit. Their agility in leaping from rock to rock . . . 
tends greatly to their protection, as neither the wily aborigine, 
nor their still greater enemy the Dingo, can follow them to their 
retreats . . . . " 

Gould writes later (1863, vol. 2, pp. 46-47) : 

"Those portions of the mountain ranges stretching along the 
eastern coast from Port Philip to Moreton Bay . . . are among the 
localities in which it is found; hills of a lower elevation than those 
of the great ranges, and the precipitous stony gullies between the 
mountains and the sea, are also situations it inhabits. . . . 

"Of its flesh as an article of food I can speak most highly, having 
frequently partaken of it in the bush and always found it excellent." 

Le Souef calls attention (1923, p. 110) to the special need of 
protective measures, since this species is entirely confined to the 
fox area of eastern Australia. He adds that in New South Wales "a 
few are found round Jenolan Caves, and at the head of the Murray 
River." He also writes (1924, p. 272) : "The Brush-tailed Rock 
Wallaby has become very scarce within Reynard's range during the 
past few years. Skins of this species used to come into the sale 
rooms in bales, now it is rare to see one." 

Musgrave writes (1925, p. 210) of the species along the Nepean 
River in New South Wales: "In former times the Brush-tailed Rock 
Wallaby . . . occurred abundantly along the banks of the river, but 
they have been so reduced in numbers by sportsmen that now they 
are but rarely seen, and it is only a matter of time before the species 
entirely disappears from the district." 


"For all their agility . . . the rock-wallabies fall victims to very 
sluggish enemies, for invariably the wallaby rocks are inhabited by 
large carpet-snakes (Python varius) , which generally lie in wait 
for their victims in the caves in which they take shelter" (Le Souef 
and Burrell, 1926, p. 202). 

Barry (1928, p. 163) reports a few in Kuringai Chase, near 
Sydney, where, "despite protective laws, shooters, foxes and hounds 
leave little chance of survival." 

"The species should survive in the more rugged or inaccessible 
parts of the Great Dividing Range in N. S. W., especially in some 
of the sanctuaries recently declared, provided such are controlled, 
and the public informed of dangers to survival of rarities, etc." 
(E. Le G. Troughton, in litt., April 16, 1937) . 

In Victoria it never occurred in great numbers and is now prob- 
ably extinct. The last record was in 1905. (C. W. Brazenor, in litt., 
March 3, 1937.) 

The species has been acclimatized on Kawau Island, New Zealand, 
where it was introduced about 1870 and now exhibits some alteration 
of coloration (Le Souef, 1930, p. 111). 

[Petrogale herberti Thomas is treated as a subspecies of P. 'in- 
ornata by Iredale and Troughton (1934, p. 43), who give its range 
as "South Queensland (about 23 to 26 S. lat.)." However, it is 
regarded as a subspecies of P. pencillata by Finlayson (1931, p. 82) , 
who writes of its status in the Dawson Valley: "Still . . . very 
numerous and widely distributed. It is to be found in thriving 
colonies in almost every range of hills away from the large towns."] 

Yellow-footed Rock-wallaby; Bar-tailed Rock- wallaby 


Petrogale xanthopus J. E. Gray, Proc. Zool. Soc. London 1854, p. 249, pi. 39 
(Mammalia), 1855. ("Australia (Richmond River?)"; this is an erroneous 
type locality, for Thomas (1888, p. 66) lists the cotypes from "Flinder's 
Range, S. A.") 

FIGS.: Proc. Zool. Soc. London 1854, pi. 39 (Mammalia); Gould, 1855, vol. 2, 
pis. 43, 44; Royal Nat. Hist., vol. 3, p. 245, fig., 1894-95; Lucas and Le 
Souef, 1909, p. 81, fig.; Le Souef and Burrell, 1926, fig. 46. 

This largest and most striking of the Rock-wallabies has disap- 
peared from many parts of its range in southern and eastern Aus- 
tralia and is in urgent need of protection to prevent its extermination. 

"Pale brown, minutely grizzled; chin and beneath white; streak 
on side from back of shoulder, and along the side of the face under 
the eye, whitish; dorsal streak narrow, brown; legs, feet, and tail 
bright yellow; end of tail more bushy and varied with brown" (J. E. 
Gray, 1855, p. 249). Fur long, soft, and silky; back of ears dark 


yellow; a brown blotch behind the elbow; a white patch on thigh 
near knee; tail more or less annulated (Thomas, 1888, p. 65). Head 
and body, 650-800 mm. ; tail, 600-650 mm. (Jones, 1924, p. 226) . 

This animal's former range included southern and eastern South 
Australia and the interior of New South Wales. It has also been 
reported from Victoria and western Queensland. 

Jones (1924, pp. 225-227) writes: 

P. xanthopus inhabits the rocky country from the Gawler Ranges to the 
Flinders Ranges, and to the eastern boundary of the State [South Australia] 
at Bimbowrie and Cockburn. . . . 

The Yellow-footed Rock Wallaby is still fairly abundant in certain parts 
of South Australia, but from many of its old haunts it has completely dis- 
appeared. It would seem that at the present time it is being driven mainly 
to the north and east of the State, and that its last stronghold in South 
Australia will be upon the New South Wales border. If it has not altogether 
disappeared from the Gawler Ranges it must now be a very rare animal, 
and in many parts of the Flinders Ranges its numbers are considerably 
reduced. From the eastern portion of the State it will almost certainly 
disappear before very many years are past, since its pelt is far too attractive 
to permit it to survive as long as the fur trade exists. Although a totally 
protected animal in this State, this protection is not extended to it by 
certain of the States upon the borders of which it lives. It is therefore not 
to be wondered at that pelts of the animal are disposed of in the markets of 
States other than South Australia, even though the animal was obtained 
within the geographical boundaries of our own State. 

Petrogale xanthopus is a fitting example of an animal which needs sanc- 
tuary for its preservation and more stringent legislative efforts to check its 

Half a century ago Lydekker wrote (1894, p. 48) : "Some hun- 
dreds of skins are annually imported to London from Adelaide, 
their value ranging from one-and-fourpence each. The skins of the 
common Rock- Wallaby [P. pencillata] are less valuable, averaging 
from threepence to ninepence each, although they have been known 
to reach as much as one-and-threepence." 

"The [Melbourne] museum has no Victorian record of this species 
though its range has been extended to the State in literature" (C. W. 
Brazenor, in litt., March 3, 1937) . 

A. S. Le Souef writes (in litt., February 15, 1937) that it is "very 
scarce, probably nearing extinction owing to settlement and the fox." 

"It provides an example of the need for unified control over State 
protection, as pelts are sold in other States though the beautiful and 
harmless marsupial is totally protected in South Australia. Such 
action may cause extermination as the animal is being driven north- 
east, and the hope of survival may rest with the sub-species de- 
scribed from south-western Queensland." (E. Le G. Troughton, in 
litt., April 16, 1937.) 

[Petrogale celeris Le Souef, described from the vicinity of Ada- 
vale, Bulloo River, southwestern Queensland, is regarded by Iredale 


and Troughton (1934, p. 44) as a subspecies of P. xanthopus. Prac- 
tically no information is at hand concerning its numerical status, 
which, however, is presumably more satisfactory than that of P. x. 

Red-necked Pademelon; Pademelon Wallaby 


Hnlmaturus Thetis "Busseuil" Lesson, Manuel Mammalogie, p. 229, 1827. 

("Port- Jackson" [Sidney], New South Wales.) 
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mamm., vol. 6, pi. 225, 1824; Lesson, 

in Bougainville, Jour. Navigation Globe Thetis et Esperance, atlas, pi. 37, 

1837; Gould, 1842, pi. 21; Gould, 1857, vol. 2, pis. 31, 32; Cabrera, 1919, 

pi. 15, fig. 5; Le Souef and Burrell, 1926, fig. 42. 

This species, formerly occurring from southern Queensland to 
Victoria, is now extinct in Victoria, and its range in New South 
Wales has become restricted to the north coast. Little seems to be 
known of its present status in Queensland. 

The upper lip is little developed, not hiding the front teeth. The 
general color above is grizzled gray; rufous on neck, shoulders, and 
rarely on cheeks and round .base of ears; a faint white hip-stripe 
sometimes present; arms and legs gray or rufous; hands and feet 
pale brown; tail gray basally, then brown above and white below. 
Head and body, 540 (female) to 660 mm. (male). (Thomas, 1888, 
p. 53.) Tail, 368 (female) to 476 mm. (male) (Waterhouse, 1846, 
vol. 1, p. 148). 

Of the early abundance of this pademelon, Gould writes (1863, 
vol. 2, p. 38) : 

Of the smaller species of Wallaby inhabiting New South Wales, the present 
is perhaps the one best known to the colonists, inasmuch as it is more 
abundant than any other. . . . All the brushes I have visited from Illawarra 
to the Hunter, as well as those of the great range which stretches along 
parallel With the coast, are equally favoured with its presence; I have also 
received specimens from Moreton Bay. . . . 

As an article of food, few animals are so valuable, its flesh being tender 
and well-flavoured, and more like that of the common Hare than that of 
any other European animal I can compare it with. 

Le Souef and Burrell remark (1926, p. 196) that it "has been 
noted in the scrub on the Blue Mountains, New South Wales." 

Troughton states (1932, p. 188) that it shows "a continual shrink- 
age in range" and is "now confined to the North Coast" of New 
South Wales. 

Lewis (1931, p. 120) apparently refers to the present species in 
the following remarks on a Victorian animal: "The Dwarf Wallaby, 
commonly known as the Paddymelon, was very plentiful once along 
the coast between Lake Wellington and Metung, but I was afraid 


that these had been exterminated. I have ascertained, however, 
that there are still some of them in this district." 

In discussing the status of this and two other species in Victoria, 
Jones and Manson say (1935, p. 35) : "All the small Wallabies are 
rare and very infrequently seen." 

More recently David H. Fleay (in Hit., June 1, 1937) reports 
the species as extinct in Victoria but as still found in New South 

Some of the older works (e, g., Thomas, 1888, p. 53; Ogilby, 1892, 
p. 53) record it from southern Queensland, and more recently Long- 
man (1930, p. 58) lists it from southeastern Queensland. However, 
Iredale and Troughton (1934, p. 46) give its current range as merely 
"New South Wales." 

Parma Wallaby or Pademelon; White-throated Wallaby or 



Hal [maturusl Parma "Gould" J. E. Gray, in Grey, Two Expeditions Discovery 
Australia, vol. 2, appendix, p. 403, 1841. ("Sidney, and its neighbourhood, 
New South Wales.") 

Fia: Gould, 1856, vol. 2, pi. 28. 

This species of New South Wales is evidently extinct. 

The general color is deep reddish brown, penciled with white and 
black; paler on sides; nape, shoulders, and forelegs brownish rust 
color; a narrow black stripe along back of neck; throat and chest 
white, rest of under parts dirty rusty white; tail scantily haired, 
black above, dirty white below (Waterhouse, 1846, vol. 1, pp. 150- 
151). Head and body, 590 (female) to 640 mm. (male) ; tail, 410 
(female) to 430 mm. (male) (Thomas, 1888, p. 58). 

Gould states (1863, vol. 2, p. 34) that in the Illawarra district 
of New South Wales "I myself saw it in a state of nature. In these 
extensive brushes it doubtless still exists, as since my return other 
specimens have been sent to me from thence by the late Mr. Strange. 
How far its range may extend westwardly towards Port Philip, or 
eastwardly in the direction of Moreton Bay, I am unable to state." 
Gould also speaks of its being hunted by the aborigines. 

"This species seems to be very rare and locally distributed" in 
New South Wales (Lydekker, 1894, p. 40). 

"The White-throated Wallaby . . . once plentiful in the Illa- 
warra district south of Sydney is apparently quite extinct, and rep- 
resented by only five specimens of which three are in England and 
two are in the Australian Museum." The uncertainty of range, as 
expressed by Gould, "will never be cleared up now, as the last of the 
two Australian Museum specimens was collected in 1889, and there 



have been no recent evidences of its possible survival." (Troughton, 
1932, p. 188.) 

The range and status of the species are given by Iredale and 
Troughton (1934, p. 46) as "New South Wales (south coast, possibly 

A. S. Le Souef remarks (in Hit., February 15, 1937) : "I think 
that this species is definitely extinct, though there may be a few 
in the dense bush near Jervis Bay." He adds that recent search 
and inquiry failed to reveal any trace of it. 

FIG. 16. Parma Wallaby or Pademelon (Thylogale parma). After Gould. 

Concerning the smaller wallabies in general, Le Souef and Burrell 
say (1926, p. 195) : "The fur is fine and soft, and great numbers 
are used for rugs, coats, and trimmings. We have practically no 
knowledge as to the individual life-histories of this group." 

Flinders Island Wallaby; Flinders Island Pademelon 


Thylogale flindersi Jones, Mammals South Australia, pt. 2, p. 240, 1924 (cf. 
Harper, 1940, p. 191). ("Flinders Island, . . . Investigator group, . . . 
Great Australian Bight.") 

This wallaby is confined to Flinders Island. "The colony was 
estimated at a hundred or so in 1924, and in view of the presence 
of rabbits as food destroyers, and cats, extinction seems certain 
unless special measures are taken. This illustrates the need for 
unsettled islands as sanctuaries, unless very large." (E. Le G. 
Troughton, in Hit., April 16, 1937.) 

The general color is grizzled light gray; sides and back of neck 
and shoulders bright rufous in the male, tawny in the female; a 


well-marked pale area along the upper lip to beneath the eye; a 
dark middorsal stripe from occiput backwards, very pronounced 
in the male; chin and throat grayish white; lower neck, chest, and 
belly colored almost like back, but somewhat lighter; limbs pale 
fawn; tail pale gray. Head and body, 510 (female) to 570 mm. 
(male) ; tail, 340 (female) to 410 mm. (male) . (Jones, 1924, p. 241.) 
Jones gives the following account (1924, p. 242) : 

Flinders observed this animal in 1802, and he records that on the island "a 
small species of Kangaroo, not bigger than a cat, was rather numerous. I 
shot five of them, and some others were killed by the botanists and their 
attendants and found to be in tolerably good condition." Even comparatively 
recently the animal was very numerous, and it has been reported that as 
many as thirty thousand were killed on the island. In 1910 a destructive 
bush fire swept the portion of the island occupied by a wallabies, and when 
I visited the place in 1920 no traces of it were to be found, and the tenant 
of the island believed it to be extinct. In 1922 I again visited the island and 
found obvious evidences of its presence, but no actual specimen was seen. 
In 1924 the little colony had considerably increased, and two specimens were 
secured for study purposes. The present small colony of wallabies occupies 
only a very limited area upon which the native bush has not been destroyed 
by various attempts at cultivation. Although the colony probably contains 
a hundred or so individuals its hold on life cannot be considered a very 
secure one. It is always at the mercy of bush fires, having no line of retreat, 
since it lives on a corner of the island that is girt by high and inaccessible 
cliffs. Moreover, it has to contend against two introduced animals, the feral 
domestic cat, which has overrun the island, and the food-destroying rabbit. 
It may at any time, though fortunately this does not seem to be at present 
the case, have to contend against human enemies. ... On account of its 
build being rather more elegant than that of the thickset Kangaroo Island 
wallaby it was at one time a favourite with people who cared to have 
wallabies running in their grounds, but at present I believe there are no 
descendants of these animals living on the mainland. It has also been an 
inhabitant of the Zoological Gardens in Adelaide, but no specimens have 
been exhibited there for many years. A former tenant of the island has 
assured me that when the wallabies were numerous there were two distinct 
types living in the island, the one obviously that described as Thylogale 
flindersi, and the other a more rare, slender, yellow wallaby. What this 
second species was it is impossible to guess; there seem to be no traces 
of it left. 

H. H. Finlayson writes (in litt., March 20, 1937) that although 
the species is plentiful in a small area, its position is insecure. 

Scrub Wallaby; Dam a Wallaby or Pademelon 


Kangurus Eugenii Desmarest, Nouv. Diet. Hist. Nat., nouv. ed., vol. 17, p. 38, 
1817. (Based upon the "kanguroo de File Eugene," Peron and Freycinet, 
Voy. Terres Australes, vol. 2, p. 117, 1816; type locality "lie Eugene, 
Josephine Archipelago," currently known as St. Peter's Island, Nuyt's 
Archipelago, South Australia.) 

FIGS.: Gould, 1841, pi. 11, and 1859, vol. 2, pis. 29, 30 (as Halmaturus der- 
bianus); Lydekker, 1894, pi. 5; Le Souef and Burrell, 1926, fig. 43. 


The present group of wallabies has long been in a state of great 
taxonomic confusion, and part of the material necessary for elucida- 
tion is evidently no longer obtainable. Under these circumstances 
the group will be treated here as a specific unit, although attention 
may be called to the three subspecies recognized by Iredale and 
Troughton (1934, pp. 46-47) : 

Thylogale eugenii eugenii (Desmarest). Type locality as stated 

Thylogale eugenii derbiana (J. E. Gray) . Type locality not stated 
in the original description but said by Waterhouse (1846, vol. 1, 
p. 155) to be "Swan River," Western Australia. Synonym: Macropus 
gracilis Gould. 

Thylogale eugenii binoe (Gould). Type locality: 'Tort Essing- 
ton," Northern Australia. This is considered an error by Iredale and 
Troughton (1934, p. 47), who substitute "Wallaby Island, Hout- 
man's Abrolhos, West Australia" (cf, J. E. Gray, List of Specimens 
Mammalia Brit. Mus., p. 91, 1843) ; however, Thomas (1888, p. 44) 
lists the type from "Port Essington, N. T. (Sir J. Richardson) 1 ' 
and places binoe in the synonymy of Macropus agilis (Gould), as 
Gould himself had already done (1863, vol. 2, p. 31). Synonyms: 
Halmaturus houtmanni Gould; H. dama Gould; H. emiliae Water- 

The former range of the species as a whole included South Aus- 
tralia, the coastal areas of southern and southwestern Western 
Australia, and various islands along the coast, including Kangaroo 
Island and Nuyt's Archipelago, South Australia, and the Recherche 
Archipelago, Garden Island, and Houtman's Abrolhos, Western Aus- 
tralia. It has become extinct on the South Australian mainland 
and on St. Peter's Island (the type locality) ; in 1910 it was reported 
as rapidly disappearing before settlement in Western Australia; 
but apparently it remains plentiful in most of its insular habitats. 

The following is adapted from Desmarest's description of what 
may be considered the paratype, which presumably came from St. 
Peter's Island: Fur soft; general color grayish brown, mixed with 
rufous near the shoulders and on the nape, crown, and forelegs; 
under parts whitish, distinctly separated from the dark color of the 
upper parts; tail grayish brown above, white below, with a slight 
reddish tint. Head and body about 21 (French) inches [567 mm.] ; 
tail, a little more than 1 (French) foot [324 mm.]. 

Gould states (1841, text to pi. 11) that he had never heard of 
"Halmaturus derbianus" being found on the mainland of South 
Australia. But he writes (1863, vol. 2, p. 36) of its abundance on 
Kangaroo Island. "The almost impenetrable scrub of dwarf Euca- 
lypti, which covers nearly the whole of Kangaroo Island, will always 
afford it a secure asylum, from which in all probability it will never 


be extirpated .... Such is the dense nature of the vegetation, that 
nothing larger than a dog can follow it; still it is taken by men 
residing on the island in the greatest abundance, both for the sake 
of its skin and its flesh: they procure it principally by snares, a 
simple noose placed on the outskirts of the brush; but they also 
shoot it when it appears on the open glades at night." 

Jones (1924, pp. 235-239) gives the following account for South 
Australia : 

Unfortunately the time has gone by when a good first-hand account of the 
small scrub wallabies inhabiting South Australia could have been written. 
The disappearance of the mainland wallabies is almost as remarkable a 
phenomenon as the disappearance of the Native Cat. . . . 

It is extremely difficult to define the former range of this complex species 
on the mainland of South Australia, or even to discriminate with any cer- 
tainty between the mainland form and the type of animal now living on 
Kangaroo Island. Only a few years ago it swarmed in scrub-covered districts 
all over the State, to-day it seems impossible to secure a single mainland 
specimen for scientific study. In places where annual battues were held by 
the present landowners less than twenty years ago it has disappeared altogether. 
It is almost certain that some still linger upon the mainland, notably at the 
southern end of Eyre's Peninsula and in the South-eastern districts, but so 
far these animals have not been properly studied or preserved. . . . 

In Kangaroo Island it is abundant and, since it lives in thousands upon 
Flinders Chase fauna reserve, it is guaranteed, in so far as complete protec- 
tion can guarantee it, perpetual survival. . . . 

The wallaby of St. Peter Island has become extinct, and therefore we 
cannot compare the animal now known as Thylogale eugenii with the St. 
Peter Island animal, and, moreover, the original specimen described by 
Desmarest is no longer in existence in Paris. It would seem to be somewhat 
doubtful if the animal now known as Thylogale eugenii is the same as the 
animal originally seen and captured on L'ile Eugene. The Kangaroo Island 
Wallaby is readily kept and bred in confinement .... 

E. Le G. Troughton remarks (in Hit., April 16, 1937) that its 
survival "appears assured on Kangaroo Island, illustrating the value 
of island sanctuaries in preserving remnants of vanishing stock." 

Hoy writes (1923, pp. 164-165) of conditions on Eyre's Peninsula, 
South Australia: "I was told, by a professional kangaroo hunter, 
that at the time of the introduction of the fox he was always sure 
of at least six dozen wallabies (Macropus eugenii) per week, but 
during the season I was there, the fourth after the introduction of 
the fox, he had not even seen one." 

According to Finlayson (1927, p. 375), "the Thylogale of the 
South Australian mainland has been exterminated before its identity 
was properly established." 

Shortridge (1910, pp. 812-813) gives its status in Western Aus- 
tralia as follows: 

"Very plentiful in many parts of the South- West, but rapidly dis- 
appearing in the cultivated districts, especially towards the northern 
end of its range. Not occurring in the coastal country between 


Albany and Cape Leeuwin, although extending to the coast at the 
Margaret River and Cape Naturaliste. Said still to exist in isolated 
patches in the North between the Swan River and Gin-Gin. Also 
occurring on the Abrolhos (Wallabi Group), Garden, and some 
of the islands off Esperance. . . . Not extending on the South Coast 
much beyond Phillips River . . . . " 

Shortridge records specimens from Arthur River, Boyadine-Dale 
River, Stockpool, Dwaladine, Woyaline Wells, Ellensbrook, and 
Twin Peak and Middle Islands, off Esperance. The accompanying 
map (p. 812) shows the former range extending in a broad coastwise 
strip from Northampton to South Australia; but the current (1910) 
range restricted to the southwestern corner of the state. 

Under the name of Macropus (Thylogale) dama, Glauert (1933, 
p. 32) gives the range of the mainland form as "South -Western 
Australia, from the Moore River in the north to the south coast 
(Cape Leeuwin and Cape Arid), inland to the Great Southern 

Troughton (1932a, p. 175) reports the species as "plentiful on the 
two largest islands" of Houtman's Abrolhos. 

Le Souef states (1930, p. Ill) that it was introduced about 1870 
on Kawau Island, New Zealand, and is still present there. 

Rufous-bellied Wallaby or Pademelon; Tasmanian Wallaby or 



Kangurus Billardierii Desmarest, Mammalogie, pt. 2, suppl., p. 512, 1822. ("La 

terre de Van-Diemen" [Tasmania].) 
FIGS.: Gould, 1841, pi. 10; Gould, I860, vol. 2, pis. 35, 36; Le Souef and Burrell, 

1926, fig. 44. 

Although this wallaby remains numerous in Tasmania and is still 
found on some of the islands of Bass Strait, it seems desirable to 
place it on record here as a vanished species of the Australian 

It is distinguished by its short ears, stout form, and long fur; 
upper parts grayish brown, tinged with olive on head and rump; 
under parts yellowish or rufous; tail short, grayish brown, the basal 
part orange above, the terminal part grayish white below (Thomas, 
1888, p. 59). Head and body, 640-765 mm.; tail, 315-320 mm. (Lord 
and Scott, 1924, p. 247). Weight, 15-20 Ib. (Gould, 1863, vol. 2, 
p. 42). 

Gould gives the following account (1863, vol. 2, p. 42) : 

I have but little doubt that the habitat of this Wallaby is limited to Van 
Diemen's Land and the larger islands in Bass's Straits, in all which localities 
it is so numerous that the thousands annually destroyed make no apparent 


diminution of its numbers. . . . Being one of the best-flavoured of the small 
Kangaroos, it is very generally eaten in Van Diemen's Land. 

The Tasmanian Wallaby may be regarded as strictly gregarious, hundreds 
generally inhabiting the same localities .... It is very easily taken with snares, 
formed of a noose placed in the run; and thousands are captured in this way, 
solely for their skins : the sportsman also may readily procure it by stationing 
himself in some open glade of limited extent, accompanied by two or three 
small yelping dogs, before which it keeps hopping round and round, and thus 
affords him an opportunity of shooting it as it passes. 


Gunn states (1838, p. 106) that "they are excellent eating, but 
the smallness of the skins renders them less valuable for tanning." 

The recent status of the Rufous-bellied Wallaby in Tasmania is 
given by Lord (1928, p. 19) : "It is evenly distributed and is plenti- 
ful in certain districts remote from settlement; but close to the 
settled areas its history is the same as the larger forms." He adds 
(p. 23) that in recent years it has increased considerably in the 
Tasmanian National Park. He also quotes (p. 24) the official 
Tasmanian returns from the hunting of this species as follows: 

1923 201,365 

1924 86,393 

1925 121,245 

1926 94,531 

Jones (1924, pp. 242-243) discusses its former occurrence in South 
Australia : 

In the collection of the British Museum there is a skull of this animal, 
formerly the property of Sir Richard Owen, which came from Mount Gambier. 
There are also two skeletons said to have been procured in South-east South 
Australia. . . . 

This is the common small wallaby of Tasmania, it is present also in some 
of the islands of Bass's Straits and on the mainland of Victoria. Evidently 
it was at one time an inhabitant of the South-eastern portion of this State, 
where the Platypus and the Koala intruded into the South Australian fauna. 
If it lingers in any corner of the South-East, I have been unable to ascertain. 
I know of no South Australian specimens. 

The species is now regarded as extinct in South Australia (David 
H. Fleay, in litt., June 1, 1937). 

Le Souef and Burrell, evidently referring to personal experience, 
say (1926, p. 196) : "M. billardieri has been met with in Gippsland 

C. W. Brazenor writes (in litt., March 3, 1937) : "Once a common 
animal in southern Victoria, the species has now entirely disap- 
peared, though it is still found on the islands of Bass Strait and in 

In the absence of the fox from Tasmania, this wallaby should 
survive indefinitely in that country. 


Whiptail, Gray-face, or Pretty-face Wallaby; Parry's Wallaby 


Macropus elegans Lambert, Trans. Linnean Soc. London, vol. 8, p. 318, pi. 16, 

1807. ("New South Wales.") 
SYNONYM: Macropus parryi Bennett. 
FIGS.: Lambert, op. cit., pi. 16; Bennett, 1835, pi. 37; Gould, 1842, pi. 19; 

Gould, 1852, vol. 2, pis. 12, 13; Lydekker, 1894, pi. 4; Le Souef and Burrell, 

1926, fig. 38. 

This wallaby, well named elegans, is rapidly diminishing in num- 
bers in its somewhat limited range in New South Wales and Queens- 
land, and is in distinct need of total protection. 

It is characterized by a slender and graceful build and a very 
long tail. The general color is clear gray, with a bluish tinge; top 
of muzzle brown, sides darker; white cheek-stripe sharply defined, 
bordered below by a gray band; ears brown at base and tip, with 
an intervening white area; digits of hands and feet black; under 
parts grayish white; tail pale gray, with a black or gray crest below 
the tip. (Thomas, 1888, p. 39.) Head and body, 732 (female) to 
793 mm. (male) ; tail, 858 (female) to 1,077 mm. (male) (Finlayson, 
19316, p. 77). 

"With this animal neither the colonists of New South Wales nor 
the naturalists of Europe are very familiar; not so much in con- 
sequence of its being really scarce, as from the extreme shyness of 
its disposition, the fleetness with which it escapes from its pursuers, 
and the mountainous and almost inaccessible parts of the country it 
inhabits. I did not succeed in procuring it myself while in Australia, 
it being confined, as far as I could learn, to the range of hills which 
stretch along parallel to the coast from Port Stephens [New South 
Wales] to Moreton Bay [Queensland], a part of the country not 
visited by me. Like most other members of its race, it is easily 
tamed, readily becoming familiar and docile." (Gould, 1842, text to 
pi. 19.) 

"Mr. Strange informs me that it inhabits the rocky ranges of the 
Clarence [New South Wales], occasionally descending into the more 
open broken country, where it frequents the ledges of rocks at an 
elevation of 2000 feet .... So fleet is this animal, that it is only 
with the assistance of the finest dogs that there is any chance of 
procuring examples ; it surpasses in fact every other animal in speed, 
and when fairly on the swing no dog can catch it." (Gould, 1863, 
vol. 2, p. 18.) 

Finlayson (19316, pp. 75-77) gives the following valuable account 
of the species in the Dawson Valley, Queensland: 

This magnificent species still occurs in large numbers in suitable tracts all 
over the valley, but in the northern part of the area is rapidly diminishing. 
In 1884 it was obtained by Lumholtz near Rockhampton and on Coomooboo- 


laroo, for instance, but is now quite unknown in the vicinity of the first place 
and on the second has become rare. . . . 

Typical of the whiptail habitats are the beautiful undulating upland parks 
of the broad-leaved ironbark (Eucalyptus siderophlora) .... As a character- 
istic example of this type of country might be cited the Grevillea plateau, 
where parryi [=eleaans] is still in very large numbers. . . . 

It is distinctly social in habit, and very likely truly gregarious, though it 
would take closer and more prolonged observation than I was able to give 
to determine the point. It certainly camps in rather large parties, 12 or 15 
being frequently seen lying up together, but in the late afternoon, when 
feeding begins in earnest, there is a tendency, I believe, for the larger males 
and females to go off in pairs. At Drumburle, where I watched it most, they 
were so numerous, however, that towards evening whole hillsides were dotted 
with the members of these disbanded camps, and it was impossible to make 
out the existence of any natural grouping. Old males are always solitary, 
as in many other species. . . . 

Like so many mammals living in open country they are very curious, and 
their curiosity has earned them a reputation for stupidity amongst trappers 
and shooters. It is said by such, that in winter when large "mobs" congre- 
gate on the sunny side of the ridges, a dozen may be shot down one by one 
before the rest make up their minds to go, provided the shooter does not 
move from his position. 

Locally it is regarded as an extremely fast wallaby, but as it is not hunted 
with dogs to any extent, it is difficult to get data for comparison with other 
species. . . . 

In considering the future of this wallaby in Queensland, there are sound 
reasons for anxiety. It is true that it is still numerous over a large area, but 
no one with any knowledge of the fate of open country species elsewhere 
would maintain that it will long survive the present rate of slaughter in the 
cattle country of the Dawson. Where man is concerned its instinct for 
self-preservation is almost nil, and as its colouration and habits make it a 
most conspicuous animal at any time, its destruction is almost a mechanical 
matter. It is very probable that the scores of thousands of whiptails which 
are killed every year in coastal Queensland, represent, not the natural increase, 
as is assumed locally, but rather the natural drainage of the species from 
large areas of relatively poor feeding grounds into smaller areas which are 
more attractive to it and which will support a denser population. When the 
country is settled these "fur pockets" act as natural traps, and destruction 
which appears to be local actually affects a much wider area, indirectly. 
It is this factor of natural concentration which is largely responsible for the 
element of unexpected suddenness which often marks the extinction of mammal 
species before advancing settlement. 

M. parryi is one of the most beautiful of Australian mammals, and is one 
of the very few species which can be easily and freely observed under natural 
conditions. It is to be hoped that its value will be recognised while there 
is still time. 

E. Le G. Troughton writes (in Hit., April 16, 1937) that this 
nearest eastern ally of the extinct Toolach was once plentiful in the 
more open coastal country from north of Sydney to the Rockhamp- 
ton district of Queensland. It is becoming rare owing to the natural 
shrinkage of habitat with settlement and to destruction for "sport" 
and profit. It is more beautiful and observable than most species 
and requires total continued protection to ensure its survival. 


Toolach; Toolache; Grey's Wallaby 

WALLABIA GRBYI (Waterhouse) 

Macropus (Halmaturus) Greyi Waterhouse, Nat. Hist. Mammalia, vol. 1, 
p. 122, 1846. ("South Australia." Iredale and Troughton (1934, p. 50) 
give, as a restricted type locality, the "Coorong, fide G. F. Angas.") 

FIGS.: Gould, 1852, vol. 2, pis. 18, 19; Finlayson, 1927, pis. 16, 17. 

This very beautiful wallaby, a former inhabitant of South Aus- 
tralia, is apparently extinct in a wild state. "One or two specimens 
in the Zoo at Adelaide are supposed to be the last living specimens 
of this species" (A. S. Le Souef , in Hit., February 15, 1937) . 

General color pale ashy brown, tinted with yellowish; under parts, 
legs, and feet pale buff-yellow; toes black; head gray; a pale yellow 
cheek-stripe, bordered above with blackish and below with brownish ; 
back of neck and back of ears pale rufous; tip of ears black; tail 
very pale gray, brown-white beneath, and with a terminal crest of 
dirty yellowish hairs. Head and body, 761 mm.; tail, 660 mm. 
(Waterhouse, 1846, vol. 1, pp. 122-124.) Additional characters given 
by Jones (1924, pp. 244-245) are: a white patch above eye; back 
with 10 to 12 dark gray bands; an ill-defined pale hip-bar. Head 
and body, 810-840 mm. ; tail, 710-730 mm. 

From the time of its discovery this species seems to have been 
almost entirely confined to southeastern South Australia, chiefly 
between the Murray River and Victoria. "Both species [Wallabia r. 
rufogrisea and W. greyi] appeared to have crossed the Murray, but 
the extent of their tenure of the river flats is difficult to estimate, 
and from this north-western part of their range they were early 
driven, or greatly reduced, by the rapid advance of closer settlement. 
Their former presence in the lower part of the county of Sturt is 
vouched for by many residents of that part of the country still 
living." (Finlayson, 1927, p. 364.) 

"The species was not exclusively confined to South Australia, but 
occurred also through a small strip of Victorian territory contiguous 
to the border" (Finlayson, 1927, p. 366). 

"Mr. Strange informs me that he met with this animal 'between 
Lake Albert and the Glenelg. The kind of country in which it is 
found consists of large open plains intersected by extensive salt 
lagoons and bordered by pine ridges. ... I never saw anything 
so swift of foot as this species: it does not appear to hurry itself 
until the dogs have got pretty close, when it bounds away like 
an antelope, with first a short jump and then a long one, leaving the 
dogs far behind it. ... I have had twenty runs in a day with four 
swift dogs and not succeeded in getting one.' " (Gould, 1863, vol. 2, 
p. 25.) 

"Many people can remember the time when Toolaches swarmed 



in the neighbourhood of Kingston. Being by far the fleetest of all 
the wallabies, its chase was at one time a very popular form of 
sport, and its beautiful pelts have been marketed in very large 
numbers in the salesrooms of Melbourne. ... It is not correct to 
say that this very fine and distinctly South Australian wallaby is 
extinct, for at the present moment five or six individuals still exist. 
Any effort to preserve this remnant must be made immediately and 
with vigour if it is to be of any service whatever." (Jones, 1924, 
p. 245.) 

FIG. 17. Toolach (Wallabia greyi). From photo. 

Finlayson furnishes an extensive account (1927, pp. 367-369), 
from which the following excerpts are taken: 

The Toolach ... in all parts of its range showed a marked partiality for 
grass country .... In the typical desert country of the counties of Russell 
and Buccleuch, where grass flats are few and far between, it occurred but 
sparsely, and here appeared to be comparatively solitary, but in the lower 
south-east, where richer soils permit a far greater development of grasses, 
its undoubted instinct towards gregariousness asserted itself, and when the 
country was first settled it was here established in a series of isolated colonies 
.... The groups . . . showed marked partiality for certain quite restricted 
areas, from which they were only driven by persistent persecution, and to 
which they returned again and again. . . . 

A considerable weight of evidence inclines me to the belief that in point 
of numbers M. greyi fell far short of the four other species of Macropus in 
the district. Although human persecution and the occupation of its chosen 


country early reduced its numbers and broke up and dispersed its larger 
colonies, it was still far from uncommon even as late as 1910, and scattered 
bands were still to be found in suitable localities. The chief of these were 
along the edges of the long strip of grass country extending from a little 
north of Millicent to the vicinity of Bull Island and Reedy Creek, and 
known locally as Avenue Valley, on the Biscuit Flat between Robe and 
Kingston, the Mosquito Plain between Naracoorte and Penola, and in the 
country between Clay Wells and Conmurra, and probably also in the sand- 
hill country of its northern district. Its rapid disappearance in the last 
twenty years may be attributed with some confidence to the invasion and 
enormous increase of the English fox, which has been proved without doubt 
to take a heavy toll of the young, even of the large kangaroos, and indeed in 
the almost unoccupied desert country where man has had little influence on 
its destinies, it seems that the fox has been the sole factor in effecting its 
extermination. Its chief natural enemies [sic] before the advent of the white 
man and the fox seems to have been the wedge-tailed eagle (Uroaetus audax), 
which, like the latter, chiefly attacked the young. These attacks were by no 
means always successful, and were sometimes thwarted by the courage of the 
females. . . . 

I learn from another source that small boys in a certain district were in 
the habit of periodically visiting the sites of eagles' nests to recover the 
scalps from the remains of young toolaches to be found lying underneath; 
this at a time when a bonus of sixpence was paid on all marsupial scalps. 

By 1923 the species had become exceedingly rare. Isolated pairs were no 
doubt scattered through the rougher stringy-bark country, but the sole 
remnant of the Toolach population which continued living in country and 
under circumstances which might be regarded as typical of that formerly 
obtaining, was a small band of perhaps fourteen individuals, located on the 
south end of Konetta sheep run, some twenty-six miles south-east of Robe. 

Public attention was first called to the rapidly approaching extinction of 
the Toolach by Professor Wood Jones, who repeatedly stressed the urgent 
need for rigid protection of this group at Konetta. In May, 1923, as there 
appeared little prospect for effective conservation in the south-east, an organ- 
ised attempt was made on a considerable scale to capture living specimens 
for transference to the sanctuary on Kangaroo Island. This, and a later 
attempt in 1924, failed in their main objective, since as a result of overmuch 
driving the four examples obtained were either dead or died shortly after 
capture, but were not altogether fruitless, as much-needed Museum material 
was thus acquired. 

The subsequent history of the species consists of a resumption of the 
exterminating process. Owing to the extensive publicity given to the two 
expeditions noted above, local attention was focussed on the Toolach to a 
degree hitherto unknown. Much of this attention was sympathetic to the 
idea of conservation, but the realization of the great rarity of the wallaby 
roused the cupidity of an unscrupulous few> and that survivors of the 1924 
attempt have been wantonly killed for the sake of the pelt as a trophy, 
is an assertion based on the admission of at least one of the slayers. The con- 
stant hunting of foxes with dogs over the Toolach country has been made the 
excuse for some of this killing, the plea being advanced that it is impossible 
to prevent the dogs running anything and everything that is put up. ... 
Interrogation usually elicits the fact that "nothing spoils a dog like checking 
him." This peculiar solicitude for the dog's training has borne very heavily 
on the Toolach and still bears very heavily on his cousin the brusher. Occa- 
sionally, however, a better spirit prevails, and recently a Toolach doe was 
promptly rescued from two kangaroo dogs which had seized her, and, in the 


patient care of Mr. J. Brown, of Robe, she has survived the rough handling 
received. She may well represent the last of her race in this State, as a 
careful and extended examination of the beat of the Konetta band by the 
writer in February of this year failed to reveal any recent traces, either 
in the shape of tracks or dejecta, and the opinion is expressed by the resi- 
dent who knows the country best that the band has been entirely extirpated. 
The species is very poorly represented in Museums, and enquiries recently 
instituted in all the States indicate that there are six skins and seven skulls 
in the public collections of Australia. 

Mr. Finlayson writes (in litt., March 20, 1937) that the species 
is nearly extinct. 

"This beautiful species presents the most tragic, and probably 
prophetic, history of all the kangaroos since white settlement" (E. 
Le G. Troughton, in litt., April 16, 1937). "The sum total of the 
isolated protective effort apparently is a doe, rescued from kangaroo 
dogs, which by now may represent the sole survivor of the species" 
(Troughton, 1938, p. 407). 

Black-gloved Wallaby; Western Brush Wallaby 


Halmaturus irma Jourdan, C. R. Acad. Sci. [Paris], vol. 5, p. 523, 1837. ("Les 
bords de la riviere des Cygnes, sur les cotes de Leuwin (Australasie)"n: 
Swan River, Western Australia.) 

SYNONYM: Macropus (Halmaturus) manicatus Gould (1841). 

FIGS.: Gould, 1841, pi. 9, and 1852, vol. 2, pis. 20, 21; Le Souef and Burrell, 
1926, fig. 39. 

Though apparently remaining common for the present in its re- 
stricted range in Western Australia, this species "requires observa- 
tion and close protection wherever possible" (E. Le G. Troughton, 
in litt., April 16, 1937) . 

Head gray above; cheeks and lips yellowish white; black spot 
under chin; back of ears brown; inside of ears yellow, the terminal 
third black; crown brown; chest, neck, sides, and outer surface of 
limbs light tawny-yellow ; wrists and tarsi yellow ; digits brown and 
black; tail with a dorsal and ventral crest, mostly gray, blackish 
toward the end, and tipped with white hairs. Head and body, 720 
mm.; tail, 630 mm. (Jourdan, 1837, p. 523.) According to Thomas 
(1888, p. 41), the general color is dark bluish gray; back of ears, 
crown, and digits black. 

"To what extent this pretty animal ranges over Western Australia 
has not been ascertained, but we know that it is very generally dif- 
fused over every part of the colony of Swan River, wherever sterile 
and scrubby districts interspersed with belts of dwarf Eucalypti 
exist .... 

"Mr. Gilbert informs us that it may be ranked among the fleetest 
of its race; that it requires dogs of the highest breed to capture it, 


and that a full-grown male weighs nearly twenty pounds. The flesh 
forms an excellent viand for the table, and the skins manufactured 
into rugs are extensively used by those whose avocations and mode 
of life lead them to spend much of their time in the bush." (Gould, 
1863, vol. 2, p. 27.) 

Thomas (1888, pp. 41-42) lists specimens from Swan River, 
Toodyay, and Perth. 

Shortridge gives the following account (1910, pp. 809-811, map) : 

Range almost identical with that of Macropus giganteus [= ocydromus], 
except that it does not seem to occur in the southern coastal districts between 
Cape Naturaliste and the Leeuwin. . . . Generally considered to be the 
best sporting animal in Western Australia. 

Not apparently dying out or disappearing even in the more thickly populated 
districts to the same extent as the smaller marsupials. 

Extending northwards beyond Watheroo, its range probably ends at some 
point to the south of Geraldton. 

Shortridge also records specimens from King River, Mount Barker, 
Boyadine-Dale River, Stockpool, Dwaladine, and Woy aline Wells. 
His map shows the range extending through the southwestern corner 
of Western Australia between Geraldton and Esperance. 

Le Souef and Burrell state (1926, p. 190): "The black-gloved 
wallaby is still very numerous in South-west Australia. . . . This 
species, strangely enough, is difficult to keep in captivity ; evidently 
it requires special food to keep it in health." They also remark 
(pp. 188-189) that "all wallabies live in or about scrub or brushwood, 
for they have three enemies ever on the look-out for stray animals 
that venture into the open, namely, the dingo, fox, and the great 
wedge-tailed eagle." 

According to Glauert (1933, p. 32) , the range of the present species 
is "South-Western Australia, from the vicinity of Geraldton in the 
north to the south coast. The eastern limit is approximately the 
No. 3 Rabbit-proof Fence. Still common near Perth." 

E. Le G. Troughton calls attention (in litt., April 16, 1937) to the 
fact that this wallaby is the nearest relative of the practically extinct 
Toolach, and suggests that "shrinkage of range should be watched 
for, although survival seems at present assured by reservations, in 
the event of more intensive cultivation." 

Tasmanian Kangaroo; Forester Kangaroo; "Boomer" 


Macropus giganteus tasmaniensis Le Souef, Australian Zool., vol. 3, pt. 4, 
p. 145, 1923. (Tasmania.) 

Formerly common and ranging practically throughout Tasmania, 
this fine species has been almost exterminated. 


Similar in size and appearance to M. giganteus, but with some- 
what coarser fur and showing greater variation in color ; upper parts 
sooty, sooty gray, rusty brown, or rusty red as a rule rusty brown, 
with grayish under parts. Head and body, 1230-1400 mm.; tail, 
970-1000 mm. (Le Souef, 19236, pp. 145, 147.) Size very large, 
form slender and graceful; tail gray, with terminal portion black; 
male about one-eighth larger than female. (Lord and Scott, 1924, 
p. 244). 

"In Van Diemen's Land, among other places, it resorts to the 
bleak, wet, and frequently snow-capped summit of Mount Welling- 
ton." In this country it "forms an object of chase, and like the 
Deer and Fox in England, is hunted with hounds; and twice a week, 
during the season, the Nimrods of this distant land may be seen, 
mounted on their fleet steeds, crossing the ferry of the Derwent, at 
Hobart Town, on their way to the hunting-ground, where they 
seldom meet without 'finding'." (Gould, 1863, vol. 2, pp. 2-3.) The 
same author gives (p. 4) an account of a hunt for an old male 
"Boomer" that led the hounds a chase of 18 miles on land, and then 
swam in the sea for more than 2 miles before he became exhausted 
and was killed. He also (p. 2) quotes R. C. Gunn to the effect that 
while the species may be found in numbers at certain places where 
food is abundant, yet it is not as a general rule gregarious and does 
not travel from place to place in flocks. 

Lord (1928, p. 18) gives the following account: 

The Forester Kangaroo formerly roamed over the greater part of Tasmania 
where conditions were suitable. It frequents, as a rule, more open country 
than M. ruficollis, and this fact, together with its larger size, is undoubtedly 
responsible for its decline. At the present time this species is met with only 
in a few localities in Tasmania. In some instances, the owners of large estates 
have taken an interest in the animal, and it is owing to the protection thus 
received that groups of this species exist to-day in certain places in the island. 

In other parts where there are scattered mobs, such as in the extreme 
North-East of Tasmania, the advance of settlement is having its effect, 
for although the species is totally protected by law, the fact must be recog- 
nised that in the more distant country districts it is a matter of extreme 
difficulty to enforce the game laws. 

Although very much reduced in numbers the Forester Kangaroo does not 
appear to be in any immediate danger of extinction, particularly if the 
landowners who have protected it in the past continue to recognise the 
variety as one worthy of being retained. Again, the species will probably be 
bred in local zoological gardens, and there is still the further possibility of this 
and other species being bred on a large scale and made an item of great 
economic importance to the State. 

More recently R. Boswell writes (in litt., May 13, 1937) that 
the species, though still wholly protected by law, has now been almost 
exterminated through excessive hunting. There has been economic 
exploitation of its hide and flesh. 


Order INSECTIVORA: Insectivores 

Family SORICIDAE: Shrews 

This family is nearly cosmopolitan in distribution but is absent 
in the Australian region (including New Guinea). There are about 
25 genera and several hundred species and subspecies. They are 
animals of generally small size and secretive habits. A single form 
is considered extinct. 

Christinas Island Musk-shrew 


Crocidura juliginosa, var. trichura Dobson, in Thomas, Proc. Zool. Soc. Lon- 
don 1888, p. 532, 1889. (Christmas Island, eastern Indian Ocean.) 

This is described as a small shrew, with skull and teeth closely 
like those of the species C. fuliginosa of the mainland (Assam, 
Tenasserim, and Malay Peninsula) . Dobson differentiated it mainly 
on the basis of a longer tail, beset with long fine hairs, but later 
collections showed that the tail as recorded by Dobson for his speci- 
men (80 mm.) was much longer than the average of 10 other speci- 
mens for which C. W. Andrews (1900) gives measurements. The 
color is not described but is doubtless, like that of the related form, 
of a dark gray. According to Andrews, the well-haired tail is the 
best character. Measurements: head and body, 65-82 mm.; tail, 
63-75 mm.; hind foot, 13-17 mm. 

The original specimen was brought back to the British Museum 
by the surveying-ship Flying-fish under command of Captain 
Maclear in 1886. Later, in 1897, a number of additional specimens 
were secured by Andrews (1900), who lists measurements in his 
Monograph of Christmas Island, and remarks: "This little animal is 
extremely common all over the island, and at night its shrill squeak, 
like the cry of a bat, can be heard on all sides. It lives in holes in 
rocks and roots of trees, and seems to feed mainly on small beetles." 
In 1908, Dr. Andrews again visited the island, to see what changes 
had taken place with the establishment of a settlement, clearing, 
and agriculture, since 1897. He found (1909, p. 102) that the shrew 
"is probably also extinct, at least no specimen was either seen or 
heard during my visit." He implies that this may have been due in 
part to cats, which had been introduced and had become numerous. 
However, cats would seem hardly sufficient to account for the ex- 
termination of a shrew, which they will kill but seldom care to eat. 
It may be that agricultural use of the land has reduced the numbers 
of the shrew about the settlement at Flyingfish Cove; nevertheless a 
careful search with modern collecting methods might still reveal the 
animal's presence. On the other hand, if it is actually gone, one 


may invoke some introduced disease, which seems, as Andrews de- 
scribes, to have been the reason for the extinction of the two native 
rats, Rattus nativitatis and R. macleari (q. v.). 


Order PRIMATES: Primates 

Family LEMURIDAE: Lemurs 

This family is restricted to Madagascar and the Comoro Islands. 
Six genera and 26 species and subspecies are recognized. While some 
forms remain common, others have become greatly reduced in num- 
bers, and one is evidently extinct. Owing to a steady reduction in 
the forested area of Madagascar and to a certain amount of perse- 
cution by the natives, the lemurs are faced with a rather uncertain 
future. Consequently, accounts of all the forms are provided in the 
following pages. 

Miller's Dwarf Lemur 


Lemur murinus J. F. Miller, Icones Anim. et Plant., pi. 13, 1777. (Madagascar.) 
SYNONYMS: Prosimia minima Boddaert (1784) ; Lemur prehensilis Kerr (1792) ; 
Lemur pusillus E. Geoffrey (1796) ; Galago madaffascariensis E. Geoffroy- 
Saint-Hilaire (1812) ; Cheirogaleus minor E. Geoffroy-Saint-Hilaire (1812) ; 
Microcebus rufus Wagner (1839); Myscebus palmarum Lesson (1840); 
Microcebus myoxinus Peters (1852) ; Chirogalus gliroides Grandidier 
(1868); Microcebus minor griseorufus Kollmann (1911). 
FIGS.: P. Brown, New Illustrate. Zool., pi. 44, 1776; J. F. Miller, Icones 
Anim. et Plant., pi. 13, 1777; G. Shaw, Cimelia Physica, pi. 13, 1796; 
Audebert, 1800, Makis, pi. 8; E. Geoffroy-Saint-Hilaire, Ann. Mus. Hist. 
Nat. [Paris], vol. 19, pi. 10, fig. 3; Peters, Reise Mossambique, Zool., I, 
Saugethiere, pi. 3, 1852; Major, 1894, pi. 1, fig. 2; Milne Edwards, Grandi- 
dier and Filhol, 1897, pi. 259, fig. 6; Kaudern, 1915, pi. 2, fig. 3. 

The comparative abundance of this species is indicated by the 
fact that the Mission Zoologique Franco-Anglo-Americaine of 1929- 
31 secured 43 specimens (Delacour, 1932, p. 220) more than of 
any other Madagascar lemur. 

Size very small; head rounded; muzzle short and pointed; eyes 
large and brilliant; ears large and naked (Forbes, 1894, vol. 1, p. 55) . 
"Two phases, rufous brown or gray. The first has the head rusty 
brown ; orbital ring and upper lip black ; stripe between eyes and on 
nose, grayish white; upper parts of body rufous brown; dorsal line 
indistinct ; sides of body and outer side of limbs mouse gray washed 
with rufous brown ; entire under parts and inner side of limbs white 
. . . ; tail rufous brown . . . ; hands and feet gray. The other phase 
is mouse gray above, the back washed with rufous, a rufous spot 
over each eye; outer side of limbs mouse gray; entire under parts 


white; tail pale rufous." Total length, about 300 mm.; tail, 150 mm. 
(Elliot, 1913, vol. 1, p. 104.) 

Schwarz (1931, p. 403) gives the range as "the whole of S.E., 
E. [=S.], and W. Madagascar, as far north as the Betsiboka River 
.... Exact limits in central Madagascar not known, but probably 
only found in the plains. Not extending farther north than Ft. 
Dauphin on the east coast." He mentions (pp. 402-403) specimens 
from: Fort Dauphin; Ankazoabo, Bara; Ambolisatra and Itampolo 
Be, north of Tulear; Tulear; and Morondava. 

Sibree (1915, p. 243) refers to this as one of the most beautiful 
and interesting of Madagascar lemurids. It "is remarkable also for 
its large and very resplendent eyes, for the eye admits so much 
light at dusk that quite an unusual brilliance is produced." 

Three specimens were captured in 1932 in the Manampetsa Re- 
serve in the southwest (Petit, 1935, p. 474) . 

"At Tabiky [inland from Cape St. Vincent], the mouse lemur 
was apparently very common and numbers were brought in alive 
by natives. ... On November 2, 1929, fifteen specimens were 
brought to me." Remains of a Microcebus were found in a pair of 
goshawks (Astur henstii) taken near Tabiky. (Rand, 1935, p. 95.) 

Smith's Dwarf Lemur 


Cheirogaleus Smithii J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 10, p. 257, 
1842. ("Madagascar"; type locality restricted by Harper (1940, p. 192) 
to "a few miles north of Fianarantsoa, central Betsileo.") 

FIGS.: Forbes, 1894, vol. 1, pi. 6; Beddard, 1902, p. 544, fig. 260. 

This lemur was reported as tolerably abundant by Shaw in 1879 
(p. 135). Although scarcely any later information is at hand, its 
nocturnal habits and its penchant for the tops of the highest trees 
have perhaps safeguarded it from serious depletion. 

Gray's type description (1842, p. 257) is as follows: "Pale brown; 
streak up the nose and forehead, the chin and beneath paler; tail 
redder." Schwarz (1931, p. 401) distinguishes this subspecies from 
M. m. murinus as follows: "Tail not longer or shorter than head 
and body. Colour above reddish brown; an indistinct dorsal band 
sometimes present. Facial streak accompanied on both sides by a 
distinct black stripe which extends as far as but hardly beyond 
the eyes." Both body and tail are about 180 mm. in length in a 
specimen from Majunga, north of the Bay of Bombetoka, which 
Lorenz-Liburnau records (1898, p. 445) as M. myoxinus. 

Schwarz (1931, p. 403) records specimens from: Vohemar, NE. 
coast; Mananara, Bay of Antongil; Mahambo, north of Foulpointe, 
NE. coast; Anabama Forest, Lake Alaotra; Ivohimanitra, Tanala; 


Vinanitelo, SE. Betsileo; and north of Fianarantsoa, central Betsileo. 
He states the range as follows: "The whole of eastern, northern, 
and east-central Madagascar, including the plateau, as far south as 
Ft. Dauphin. Also the north-west, down to the Bay of Bombetoka." 
If, however, Schwarz is correct (p. 402) in recording murinus from 
Fort Dauphin, the range of smithii can scarcely extend quite so 
far south. 
G. A. Shaw (1879, pp. 135-136) gives the following account: 

They inhabit a belt of forest-land stretching from the eastern forest into 
the heart of Betsileo, a few miles north of Fianarantsoa, where they are 
tolerably abundant. They live on the tops of the highest trees, choosing 
invariably the smallest branches .... 

Their food consists of fruit and insects and most probably honey. I have 
frequently seen them catching the flies that have entered their cage for 
the honey; and I have supplied them with moths and butterflies, which they 
have devoured with avidity. 

They are extremely shy and wild. Although I have had between thirty and 
forty caged at different times, I have never succeeded in taming one. . . . 

I have had none breed in captivity. 

Kaudern (1915, p. 74) records several specimens (as M. minor] 
from Ste. Marie de Marovoay on the Betsiboka River in the north- 
west, and one specimen (as M. smithii) from Fenerive on the east 

G. M. Allen (1918, p. 516) records a specimen from Didy, south 
of Lake Alaotra. 

Coquerel's Dwarf Lemur 


Cheirogalus Coquereli Grandidier, Rev. Mag. Zool., ser. 2, vol. 19, p. 85, 1867. 

("Morondava," west coast of Madagascar.) 
SYNONYM: Microcebus coquereli Schlegel and Pollen (1868). 
FIGS.: Schlegel and Pollen, 1868, pi. 6; Milne Edwards, Grandidier and Filhol, 

1897, pi. 259, fig. 4; Beddard, 1902, p. 544, fig. 261; Elliot, 1913, vol. 1, 

pi. 4, upper fig. (facing p. 145). 

To judge by the small number of specimens recorded, this is one 
of the rarest lemurs of Madagascar. 

It is a little smaller than Phaner furcifer; above dark gray, washed 
with rufous ; tail dark rufous, except at the base, where it is colored 
like the back; under parts yellowish gray. Head and body, 210 mm.; 
tail, 340 mm. (Grandidier, 1867a, p. 85.) 

Grandidier (1867a, p. 85) had seven of these animals in his pos- 
session. They were nocturnal and lived on leaves and fruit. 

Schlegel and Pollen state (1868, p. 13) that the species inhabits 
the most impenetrable forests. They had only a single specimen, 
secured in the forests of Congony, inland from the Bay of Passan- 
dava, in northwestern Madagascar. 


According to Elliot (1913, vol. 1, p. 107), the range extends on 
the west coast from Cape St. Vincent to Helville, in the vicinity of 
the Bay of Passandava. 

Only five specimens are reported by Delacour (1932, p. 220) as 
collected by the Mission Zoologique Franco-Anglo-Americaine of 

Geoffrey's Fat-tailed Lemur 


Cheirogaleus medius [E.] Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris], 
vol. 19, p. 172, 1812. (Type locality not stated; restricted by Schwarz 
(1931, p. 405) to "Ft. Dauphin, S.E. Madagascar.") 

SYNONYM: Opolemur thomasi Major (1894). 

FIGS.: E. Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris], vol. 19, pi. 10, 
fig. 2, 1812; Major, 1894, pi. 1, fig. 1. 

Extremely little information is available concerning this lemur, 
but it is evidently one of the less common of the Madagascar forms. 

Head broad; snout short; upper parts gray, with a wash of rusty 
brown, the tips of the hairs silvery; a whitish band extending from 
between the eyes to the naked nose-pad; a whitish half-collar on 
each side of the neck; orbital ring and ears brownish black; under 
parts, inner side of limbs, and hands and feet yellowish white. Head 
and body of female, 232 mm.; tail, 195 mm. (Major, 1894, p. 20; 
type description of Opolemur thomasi.) 

Major (1894, p. 20) records three specimens (as 0. thomasi) from 
Fort Dauphin. "Nothing is known of the distribution . . . north of 
Ft. Dauphin in eastern Madagascar" (Schwarz, 1931, p. 405). 

The Mission Zoologique Franco-Anglo-Americaine of 1929-31 col- 
lected nine specimens (Delacour, 1932, p. 220). 

"These little lemurs are apparently entirely nocturnal .... At 
Tabiky [inland from Cape St. Vincent], I found them in a gallery 
forest through savannah and dry brush." (Rand, 1935, p. 95.) 

Neither Delacour nor Rand gave the subspecific determination of 
their specimens. 

Samat's Fat-tailed Lemur 


Chiroffalus Samati Grandidier, Rev. Mag. Zool., ser. 2, vol. 20, p. 49, 1868. 

("Flumen Tsidsibon in littore occidentali Madagascar insulae.") 
FIG.: Proc. Zool. Soc. London 1872, pi. 70. 

The few known specimens of this lemur come from a limited 
section of the west coast of Madagascar (Tsidsobon River to Mo- 
rondava) . 


Fur of body and tail rather short; dark gray above, fulvous below; 
tail fat, faded rufous; a white stripe from forehead to nose; orbital 
ring black. Head and body, 190 mm.; tail, 170 mm. (Grandidier, 
1868, p. 49.) 

Major (1894, p. 18) and Schwarz (1931, p. 405) record specimens 
from Morondava. 

Milius's Mouse Lemur 


Cheirogaleus major [E.] Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris], 

vol. 19, p. 172, 1812. (Type locality not stated; restricted by Schwarz 

(1931, p. 406) to "Fort Dauphin, S.E. Madagascar.") 
SYNONYMS: Lemur commersonii Wolf (1822); Cheirogaleus milii E. Geoffrey 

(1828); Ch. typicus A. Smith (1833); Mioxicebus griseus Lesson (1840); 

Chirogalus adipicaudatus Grandidier (1868). 
FIGS.: E. Geoffroy, Ann. Mus. Hist. Nat. [Paris], vol. 19, pi. 10, fig. 1, 1812; 

Geoffrey and Cuvier, Hist. Nat. Mamm., livr. 32, pi. 188, 1821; Proc. 

Zool. Soc. London 1879, pi. 9 (ssp.?); Milne Edwards, Grandidier and 

Filhol, 1897, pi. 259, fig. 5; Elliot, 1913, vol. 1, pi. 5 (ssp.?). 

The meager information we have concerning this lemur is an 
indication of its rarity. 

Upper parts varying from brownish gray to ashy brown; under 
parts and inner side of limbs yellowish or whitish; orbital ring 
black; nose light gray; hands and feet dark brown; tail pale rufous 
or ashy brown, sometimes with white tip. Total length, 580 mm.; 
tail, 275 mm. (Elliot, 1913, vol. 1, pp. 93-94.) Ears naked for their 
distal half; color more grayish than in Ch. m. crossleyi (Schwarz, 
1931, p. 405). 

The respective ranges of Ch. m. major and Ch. m. crossleyi are 
none too clearly defined in the available literature. The former 
seems to occur in the south and west of Madagascar; the latter, in 
the center and the northeast. 

Elliot's statement (1913, vol. 1, p. 93) of the range of the present 
form is probably not altogether accurate: "Eastern coast of Mada- 
gascar; Fort Dauphin to Tamatave; also in the lower wooded 
regions of Betsileo Province; and on the west coast from Tullare 
[=Tullear] to Pasandava, Central Madagascar." 

G. A. Shaw (1879, pp. 134-135) records a specimen (subspecies 
not determined) from the forests on the eastern side of Betsileo. "Its 
food consists of fruits and possibly honey .... It appears to be a 
very uncommon animal, ... as this is the only specimen I have 
been able to obtain, although I kept a man in the forest for two 
months seeking for one after I had obtained this one." 

Major (1894, p. 22) records specimens from Morondava in the 
southwest and from Tamatave and Ankay Forest in the northeast. 


"Milius' Mouse-Lemur, though a rare species, is widely distributed 
in Madagascar" (Forbes, 1894, p. 51). 

Kaudern (1915, p. 74) records a specimen from Ste. Marie de 
Marovoay on the Betsiboka River, northwestern Madagascar (where 
the animal is said to be rather rare), and several specimens from 
Andranolava, north central Madagascar. (Here again, the speci- 
mens do not appear to have been determined subspecifically.) 

Delacour (1932, p. 219) reports only six specimens collected by 
the Mission Zoologique Franco-Anglo-Americaine of 1929-31. 

Crossley's Mouse Lemur 


Chirogaliis crossleyi Grandidier, Rev. Mag. Zool., ser. 2, vol. 22, p. 49, 1870. 

("Forets est d'Antsianak," Madagascar.) 
SYNONYMS: Chirogale melanotis Major (1894); Chirogale sibreei Major 

FIGS.: Proc. Zool. Soc. London 1872, pi. 71, nearer fig.; Forbes, 1894, pi. 5. 

This subspecies is apparently even rarer than Ch. m. major. 

Upper parts rufous, especially on the head, under parts whitish; 
head very large, rounded; orbital ring black; inner surface of ears 
covered with dark brown hairs; tips of ears bordered with black; 
tail short and well furred. Body, 200 mm.; tail, 120 mm. (Gran- 
didier, 1870, p. 49.) Ears hairy inside and out, with hardly a naked 
tip; fore parts of body strongly washed with brownish (Schwarz, 
1931, p. 405). The tail of the type specimen of crossleyi was evi- 
dently defective; the types of "melanotis" and "sibreei" have a 
total length of 490-500 mm., and a tail length of 225-250 mm. 
(Elliot, 1913, vol. 1, pp. 95-96). 

This lemur seems to be scarcely known except from the type 
specimens of crossleyi, "melanotis" and "sibreei" These are, re- 
spectively, from the forests east of Antsianak; from Vohima on the 
northeast coast; and from Ankeramadinika, one day's journey to 
the east of Antananarivo. Thus the known range extends from east 
central to northeastern Madagascar. 

Hairy-eared Mouse Lemur; Tufted-eared Mouse Lemur 


Chirogaleus trichotis Gunther, Proc. Zool. Soc. London 1875, p. 78, 1875. (On 
the "way from Tamantave to Murundava," Madagascar.) 

FIGS.: Gunther, 18756, pi. 15, p. 79, figs. 1, 2; Milne Edwards, Grandidier and 
Filhol, 1897, pi. 259, fig. 9. 

Since the single specimen of the Hairy-eared Mouse Lemur was 
described in 1875, no subsequent specimen has turned up, despite 
the extensive collecting that has been carried on in Madagascar in 


the meantime. It seems fairly necessary, therefore, to class it 
among the extinct species. 

"Brownish grey; lower parts grey, with the hairs white-tipped. 
A triangular spot in front of the eye black; median line of the 
snout and lips whitish. Hands and feet grey, with white-tipped 
hairs. Ears very short, hidden in the fur. The lower part of the 
concha and the space before the ear covered with a tuft of very 
long hairs. Tail rather shorter than the body, covered with short 
hairs . . , . All the nails pointed, claw-like." Head and body, 152 
mm.; tail, 149 mm. (Gunther, 18756, pp. 78-79.) "This species 
differs from all the members of this genus in the tufts of hair 
standing out from the ears and sides of head, above the ears" (Elliot, 
1913, vol. l,p. 97). 

Elliot's authority for giving (p. 96) the range of the species as 
the "forests of Antsianak" is not apparent. The exact type locality 
is unknown, and no information is at hand concerning the route fol- 
lowed by Crossley, the collector of the type, on his way from Ta- 
matave on the east coast to Morondava on the west coast. 

Fork-marked Lemur. Maki a fourche (Fr.) 


L[emur] furdfer Blainville, Osteogr., Mammif., Primates, Lemur, p. 35, 1839. 

(Type locality not stated originally; "probably the region of the Bay of 

Antongil, N.E. Madagascar" (Schwarz, 1931, p. 407).) 
FIGS.: Blainville, Osteogr., Mammif., Primates, Atlas, Lemur, pi. 7, 1839; 

Schlegel and Pollen, 1868, pi. 5; Milne Edwards, Grandidier and Filhol, 

1897, pi. 259, fig. 3. 

The Fork-marked Lemur was formerly abundant in Madagascar 
and is still fairly common. 

Upper parts reddish gray ; outer side of limbs dark rufous ; throat 
pale rufous; chin and rest of under parts yellowish; a black stripe 
from lower part of back to crown, where it bifurcates, one branch 
ending over each eye; hands and feet dark brown; tail bushy, dark 
reddish brown with black tip. Total length about 600 mm.; tail, 
350 mm. (Elliot, 1913, vol. 1, p. 109.) 

"This species has been recorded by Pollen and van Dam from 
various localities on the N.W. coast, north of the Bay of Bombetoka 
(Bay of Ampasindava, Jangoa River, Kongony River), but also 
farther south at Morondava, on the W. coast. Found by M. J. 
Audebert at Tassumbe/ N.E. coast." (Schwarz, 1931, p. 407.) He 
also states that "there is no definite record of the occurrence of this 
species considerably south of the Bay of Antongil." He mentions 
having examined specimens from Vohemar, NE. coast, and An- 
doany, NW. coast. 


This pretty species is found in abundance in the forests of western 
Madagascar, and it also appears to inhabit the eastern part. The 
natives state that it is very fond of honey. It is nocturnal and its 
chase is extremely difficult. (Schlegel and Pollen, 1868, pp. 9-10.) 

Twelve specimens were collected by the Mission Zoologique 
Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) . 

"Near Tabiky [inland from Cape St. Vincent] , in November, 1929, 
I found the squirrel lemur fairly common about my camp in a gallery 
forest through savanna and low dry brush. Usually found in pairs, 
. . . they sometimes moved about rapidly through the tops of the 
tall trees, but more often were in the lower trees and bushes .... 

"In the rain forest on Mt. d'Ambre [in the extreme north] this 
lemur was fairly common, and noisy throughout the night, but it 
kept to the tops of the forest trees." (Rand, 1935, p. 95.) 

Broad-nosed Gentle Lemur 


Hapalemur simus J. E. Gray, Cat. Monkeys, Lemurs, and Fruit-eating Bats 

Brit. Mus., p. 133, 1870. ("Madagascar.") 
FIGS.: Gray, Proc. Zool. Soc. London 1870, pi. 52; Milne Edwards, Grandidier 

and Filhol, 1896, pis. 122 A, 122 D (fig. 1), 122 E. 

Comparatively few specimens of this species seem to be known, 
and the information concerning it (other than anatomical) is very 

<r Nose broad and truncated; . . . back iron-grey, with a rufous 
tinge; the hairs black, with a subapical rufous band, and the lower 
part lead-coloured; throat whitish; patch on rump at base of tail 
yellowish" (J. E. Gray, 1870, p. 133). Forbes (1894, pp. 82-83) 
adds the following details: ears short, covered with long hair; 
sides of head, neck, and region round the eyes lighter than the 
back; lower back, sides of body, and outer surface of limbs sooty 
gray, with here and there a wash of rufous; tail, except at base, 
sooty gray; under side of body and inner side of arms pale sooty 
gray; no patch of spines on the arm above the wrist as in H. griseus. 
Schwarz (1931, p. 407) gives a total length of 900 mm. 

"It would appear . . . that the distribution . . . includes the 
whole forested region of eastern Madagascar. It has not been 
recorded up to now from the north-west." Single specimens from 
Nandihizana, central Betsileo, S.E. Madagascar, and from Passum- 
bee, N.E. coast, are mentioned. (Schwarz, 1931, pp. 407-408.) 

G. A. Shaw writes (1879, pp. 133-134) of a live specimen that 
"came from the higher-level forests on the eastern side of the 
Betsileo, among the bamboos, on which it appears in a great mea- 
sure to subsist. ... I have tempted it with very many different 


kinds of berries and fruits growing in the forest; but it would not 
touch any of them." It fed steadily and regularly upon grass. 

G. M. Allen (1918, p. 516) records two specimens from near Am- 

An indication of the rarity of this species is the fact that no speci- 
mens were reported by the Mission Zoologique Franco-Anglo-Ameri- 
caine of 1929-31. 

Gray Lemur 


L[emur] griseus Link, Beytr. Naturg., vol. 1, pt. 2, p. 65, 1795. (Based upon 

"Le petit Maki gris" of Buffon (Hist. Nat., suppl., vol. 7, p. 121, 1789); 

type locality, Madagascar. ) 
SYNONYMS: Lemur griseus E. Geoffroy (1796); Lemur cinereus Desmarest 

(1820); Hapalemur schlegeli Pocock (1917). 
FIGS.: Buffon, Hist. Nat., suppl., vol. 7, pi. 34, 1789; Audebert, 1800, pi. 7; 

Schlegel and Pollen, 1868, pi. 3; Milne Edwards, Grandidier and Filhol, 

1896, pis. 122 B, 122 D (fig. 2), 122 F. 

Extremely little new information concerning this subspecies has 
come to light during the past 70 years. It must have become 
quite rare. 

Upper parts light olive-brown, brighter on top of the head; rest 
of head gray; cheeks, throat, breast, and inner side of limbs ochra- 
ceous-white; tail a little darker than the back. Total length, 24 
inches; tail, 13^ inches. (Schlegel and Pollen, 1868, p. 7; Forbes, 
1894, vol. 1, p. 81.) The general grayish green of this subspecies 
is contrasted with the reddish green of H . g. olivaceus. The presence 
of a wrist gland distinguishes both of these forms from H. simus. 
(Schwarz, 1931, p. 408.) 

Schwarz (1931, p. 408) gives the range of the present form as 
follows: "The whole south and west, and the dry central plateau 
as far east as Lake Alaotra ; it also goes north beyond the Betsiboka 
River in the north-west." He records specimens from the following 
localities: Lake Alaotra and Ambatondrazaka, central northeastern 
Madagascar; District Ambalavo, in the southeast; and Tany Ma- 
landi, in the northwest. 

According to Schlegel and Pollen (1868, pp. 7-8), this lemur in 
northwestern Madagascar inhabits by preference the forests of 
bamboo. It was found at a few days' journey from the coast, along 
the Ambassuana River in the Tanimalandy district. It is entirely 
nocturnal, and sleeps during the day on the highest stems of the 
bamboos. The stomachs of all specimens were found filled with 
bamboo leaves. 

Delacour (1932, p. 219) records 25 specimens of "Hapalemur 
griseus" as collected by the Mission Zoologique Franco-Anglo-Ameri- 


caine of 1929-31. However, the further account given by Rand 
(1935, p. 95) indicates that these represent the subspecies olivaceus. 

Olivaceous Lemur 


Hapalemur olivaceus I. Geoffroy, Cat. Method. Mamm. [Mus. Paris], pt. 1, 
Primates, p. 75, 1851. ("Madagascar." Type locality restricted by Elliot 
(1913, vol. 1, p. 127) to "Ampazenambe, Madagascar.") 

FIGS.: Forbes, 1894, vol. 1, pi. 8 (ssp.?) ; Milne Edwards, Grandidier and 
Filhol, 1896, pis. 122 C, 122 D (fig. 3). 

This subspecies appears to be considerably commoner than E.g. 

It is similar to the latter but has a longer and denser pelage; 
color olive, with a rufous tint; throat gray rather than white; 
cheeks speckled gray (I. Geoffroy, 1851, p. 75). 

According to Schwarz (1931, pp. 408-409) , this lemur "is found in 
the moist and wooded north-east and east [of Madagascar], prob- 
ably as far south as Fort Dauphin." He records specimens from 
the following localities: Mananare, Bay of Antongil; Vohemar, 
NE. coast; Tamatave, E. coast; Analamazaotra, east of Tananarive; 
Ambohimitombo, Tanala country, E. Madagascar; Vinanitelo, S. 
Betsileo ; and several localities on the east coast north of Tamatave. 

Kaudern (1915, pp. 70-71) reports the animal as apparently not 
rare in the forests west of Fenerive on the east coast, where he 
obtained three specimens from the natives. 

Rand writes (1935, p. 95) that it is diurnal. He found it in 
the tops of the lower trees in the rain forest, and occasionally in 
dense thickets of bamboo on the edge of the forest. "Occasionally 
found singly, it was more often seen in groups of two or three. At 
Manombe in the southeast I saw two running about through the 
forest tree tops .... Hapalemur was fairly common about camp 
two days northeast of Maroantsetra." 

Ring-tailed Lemur 


[Lemur] Catta Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 30, 1758. (Based upon 
the "Maucauco" of Edwards (Nat. Hist. Birds, pt. 4, p. 197, 1751); 
type locality, "Madagascar.") 

FIGS.: Edwards, 1751, pi. 197; Schreber, Saugthiere, vol. 1, pi. 41, 1774; Aude- 
bert, Hist. Nat. Makis, pi. 4, 1800; Geoffroy and Cuvier, Hist. Nat. Mamm., 
livr. 5, pi. 27, 1819; Milne Edwards and Grandidier, 1890, pis. 171, 172; 
Royal Nat. Hist., vol. 1, p. 211, fig., 1893-94; Proc. Zool. Soc. London 
1906, p. 124, fig. 48; Elliot, 1907, p. 545, fig. 76; Elliot, 1913, vol. 1, pi. 5, 
upper fig. 


This is perhaps the best known of all the Madagascar lemurs, 
and it still appears to be moderately common. 

The fur is soft and delicate; face and ears white; nose and area 
about each eye black; top and back of head dark ashy; back and 
sides of a redder ash color ; outer side of limbs light ashy ; upper sides 
of paws whitish ; under parts and inner sides of limbs white ; tail with 
broad alternate rings of black and white (Edwards, 1751, p. 197). 
"Length of body and tail together, 40 inches" (Forbes, 1894, vol. 1, 
p. 76). 

"This species, which inhabits rocky open country, is found in 
south-western, southern, arid south-eastern Madagascar" (Schwarz, 
1931, p. 410). 

"As far as my experience of seven years goes, these Lemurs are 
found only in the south and south-western borders of the Betsileo 
province of Madagascar." They are not found in the forests, but 
among the rocks. "The prickly pear . . . constitutes their chief 
article of winter food .... Their summer food consists of different 
kinds of wild figs and bananas." (G. A. Shaw, 1879, pp. 132-133.) 

This species bears captivity well, and is everywhere offered for 
sale by the natives (Kaudern, 1915, p. 50). 

Schwarz (1931, p. 410) records specimens from Tulear in the 
southwest, and adds: "It has been found by van Dam at Moron- 
dava, Matseroka, and the Bay of St. Augustin in the south-west, 
and is recorded by him as far north-east as the region of Ft. 

This species ranges over a vast area, from Mangoky on the 
north to beyond Menarandra on the south. Decary records it in 
Androy along all the rivers and as far as the region of Beloha. 
Perrier de la Bathie has observed bands on the western parts of tho 
massif of Andringitra. (Petit, 1931, p. 560.) 

Thirty specimens were collected by the Mission Zoologique Franco- 
Anglo-Americaine of 1929-31 (Delacour, 1932, p. 219) . 

Rand (1935, p. 96) mentions observations on this lemur at 
Ampotaka and Lake Tsimanampetsotsa and near Tulear. He also 
writes (pp. 95-97) : "The ring-tailed lemur was found in and about 
most of the more densely -wooded areas and the gallery forest in the 
arid parts of southwestern Madagascar. 

"It is a diurnal and crepuscular creature .... 

"This animal, like most of the lemurs, is gregarious. It was usually 
seen in parties of from four or five up to ten or fifteen and more. . . . 

"Wild specimens were seen to eat leaves of certain trees .... 

"They are often kept as pets by Europeans and are carried to 
various parts of the island." 

At the Manampetsa Reserve in the southwest numerous bands 
were seen in 1926, but the species was rare in 1933 (Petit, 1935, 
p. 474). 


Black Lemur 


[Lemur] Macaco Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 44, 1766. ("Mada- 

SYNONYM: Lemur leucomystax Bartlett (1863). 

FIGS.: Schreber, Saugthiere, vol. 1, pi. 40 A, 1774; Proc. Zool. Soc. London 
1862, pi. 41; Schlegel and Pollen, 1868, pi. 1; Proc. Zool. Soc. London 1880, 
p. 451, fig. 1; Proc. Zool. Soc. London 1885, p. 672, fig.; Milne Edwards and 
Grandidier, 1890, pis. 130-132. 

The Black Lemur still occurs in fairly large bands in the north- 
west of Madagascar. 

It has the ears tufted, with long hairs continuing down the side 
of the neck to the angle of the mouth. The male is entirely black. 
Female: snout and back of head black; forehead blackish gray; 
whiskers and ear-tufts white; general body color rich ferruginous 
brown; limbs and neck reddish yellow; tail whiter; under parts 
and inner side of limbs creamy white. There is a considerable degree 
of variation in the color of this species. (Forbes, 1894, vol. 1, p. 70.) 
Total length, 41 inches; tail, 22 inches (Schlegel, 1876, p. 303). 

"The range ... is limited to the forests of the N.W. coast, north 
of the Bay of Bombetoka and the coast islands. It has been recorded 
by Pollen and van Dam from the following places: Anorontsanga ; 
Syrangene; Kongony and Jangoa Rivers; Andoany, Narendry Bay 
("Maroandiana")." (Schwarz, 1931, p. 417.) 

These animals inhabit the forests extending between the Bay of 
Diego-Juarez and the Bay of Bombetoka, as well as the forest of 
Loucoube in the isle of Nossi-Be. They live in bands in the highest 
trees of the impenetrable forests. Bananas are their ordinary food 
in the wild state. They are also fond of the brains of birds. (Schlegel 
and Pollen, 1868, p. 2.) 

The range includes the upper Sambirano (Petit, 1931, p. 562). 

Eight specimens were collected by the Mission Zoologique Franco- 
Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) . 

"The parties observed were much larger than those of that species 
[L. julvus], containing sometimes as many as 18 individuals 
males, females, old, and young. They were always very tame." 
(Rand, 1935, p. 99.) 

Sanford's Lemur 


Lemur julvus sanjordi Archbold, Am. Mus. Novit., no. 518, p. 1, 1932. ("Mt. 
D'Ambre, Madagascar.") 

This lemur is known only from 18 specimens collected in the 
type locality by the Mission Zoologique Franco-Anglo-Americaine 
of 1929-31. 


It differs from all the black-nosed lemurs, except Lemur macaco 
rujus, in its lighter color, and from the latter in its cheek and ear 
tufts, in which it resembles Lemur macaco. Nose black; top of head 
dark olive-buff; general color of upper parts snuff-brown to drab; 
hands russet ; spot at root of tail and basal half of tail bay-colored ; 
distal half of tail bushy, the hairs with blackish brown tips and with 
a subterminal clay-colored band; a ruff of long hairs around the 
ears and down the cheeks, varying from white to light ochraceous- 
buff; under parts gull-gray, washed with buffy. Female without 
cheek-tufts. Total length of male, 895 mm.; tail, 495 mm. 

This form is restricted to the rain forest of Mount D'Ambre, 
northern Madagascar. (Archbold, 1932, p. 1.) 

White-fronted Lemur; White-faced Lemur. Maki a front 

blanc (Fr.) 


L[emur] Albifrons [E.] Geoffroy, Mag. Encycl. [2d yr.], vol. 1, p. 48, 1796. 
(Type locality not stated in original description, but later given as "Mada- 
gascar" (E. Geoffroy-Saint-Hilaire, 1812, p. 160).) 

SYNONYM: Prosimia frederici Lesson (1840). 

FIGS.: Audebert, Hist. Nat. Makis, pi. 3 (facing p. 13), 1800; Geoffroy and 
Cuvier, Hist. Nat. Mamm., livr. 3, pis. 17, 18, 1819; Milne Edwards and 
Grandidier, 1890, pis. 136, 144, 154, 155. 

To judge by the 36 specimens collected by the Mission Zoologique 
Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) , this 
must be one of the commonest of Madagascar lemurs. 

Pelage brown; top of head, cheeks, and jaws white (. Geoffroy, 
1796, p. 48). "Of this race there are two mutations which occur 
together, viz., (1) a normally black-headed, whitish-cheeked type, 
with dark ground-colour, in which the female is only slightly paler 
than the male, and (2) the 'albifrons' type, which shows a reduction 
of black pigment, the ground-colour being more reddish, especially 
so in the female; the male has the whole crown, cheeks, and beard 
white or whitish, but there are females which show a whitening on 
the head, although the normal phase of the 'albifrons' female has a 
lead-grey head and a grey muzzle" (Schwarz, 1931, p. 410). 

"The range of this local race apparently includes the north- 
eastern coast of Madagascar as far as, and slightly beyond, the 
Bay of Antongil. The northern and western limit is uncertain; 
only the dark phase apparently occurs on the north-west coast." 
Specimens are recorded from: Vohemar and Sahambavany, N.E. 
coast; Mananare, Maroansetra, Androutse, and Ampazenardo, in 
the vicinity of the Bay of Antongil. (Schwarz, 1931, pp. 410-411.) 

"The effects of hunting by natives on the animal life of Mada- 
gascar are negligible. The main destruction of the fauna is caused 


by the cutting of the forests. But near Maroantsetra we saw what 
might happen. Near the town itself these lemurs were scarce and 
very wary, apparently having been hunted with guns, but once 
away from large settlements they were very common and tame. On 
July 22, 1930, two days northwest of Maroantsetra, I surprised a 
party of eight in the low bushes of the ground cover in the forest. 
They fled but a little way and I sat down to watch them. Very soon 
the whole party came back and resumed feeding on the fruit of a 
low bush near me. 

"Parties of these lemurs were often heard grunting and growling 
in the forest as though fighting." (Rand, 1935, p. 98.) 

Brown Lemur; Fulvous Lemur. Maki brun (Fr.) 


L[emur] Fulvus [E.] Geoffroy, Mag. Encycl. [2d yr.], vol. 1, p. 47, 1796. 

("Madagascar." This subspecies is considered typified by specimens from 

the Tamatave region (Schwarz, 1931, p. 411).) 
SYNONYMS: Prosimia macromongoz Lesson (1840); Lemur bruneus van der 

Hoeven (1844). 
FIGS.: Buffon, Hist. Nat., suppl., vol. 7, pi. 33, 1789; Lacepede and Cuvier, 

1801, unnumbered pi. 

Although it is said that "this race is about the most common 
Lemur in captivity" (Schwarz, 1931, p. 412), very little information 
can be offered concerning it, owing partly to the confusion that has 
long prevailed in the taxonomy and nomenclature of this specific 

The pelage is brown above, gray below; head black; hands fulvous 
or brown (]5. Geoffroy, 1796, p. 47). The animal is a third larger 
than Lemur mongoz; its tail, less bushy and more woolly, tapers 
toward the tip; rump and legs washed with olive (fi. Geoffroy, in 
Lacepede and Cuvier, 1801, p. 3 of "Le Maki Mococo et le Maki 
brun"). Ground color olive-brown; cheeks yellowish white (Schwarz, 
1931, p. 410). 

"The range of this race is not completely known. It obviously 
inhabits the coast between the Bay of Antongil and Andovoranto, 
but may go farther south, as far as Mahanoro. In the interior it 
appears to go into the forest-belt east of Tananarive." Specimens 
are recorded from: Andragoloaka, S.E. of Tananarive, Prov. Ime- 
rina; Lakato Forest, Ankay, N.E. of Tananarive, Imerina; Sakana 
and Ambotorao, opposite the He Ste. Marie; and Tamatave. 
(Schwarz, 1931, p. 411.) 


Collared Lemur. Maki a f raise (Fr.) 

LEMUR MACACO OOLLARIS E. Geoffrey-Sain t-Hilaire 

Lemur collaris [E.] Geoffroy-Saint-Hilaire, Ann. Mus. Hist. Nat. [Paris], 

vol. 19, p. 161, 1812. ("Madagascar.") 
SYNONYMS: Prosimia melanocephala J. E. Gray (1863); P. xanthomystax 

J. E. Gray (1863); ?F. flavifrons J. E. Gray (1867); Lemur nigerrimus 

P. L. Sclater (1880); Lemur mongoz var. cinereiceps Milne Edwards and 

Grandidier (1890). 
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mamm., livr. 2, pi. 11, 1819; Proc. 

Zool. Soc. London 1863, pi. 17 (as Prosimia xanthomystax), pi. 18 (as P. 

melanocephala) ; ?Proc. Zool. Soc. London 1867, pi. 31 (as P. flavifrons) ; 

?Proc. Zool. Soc. London 1880, p. 451, fig. 2 (as Lemur nigerrimus) ; 

Milne Edwards and Grandidier, 1890, pis. 140, 147 (as L. mongoz var. 

cinereiceps); Elliot, 1913, vol. 1, pi. 6 (as L. julvus). 

This lemur appears to be common in southeastern Madagascar. 

Pelage rufous-brown above, fulvous below; a ruff of rufous 
hairs; face lead-colored (E. Geoffroy-Saint-Hilaire, 1812, p. 161). 
Cheeks orange-yellow; ground color light brown; a faint spinal 
line generally present; female generally with head gray. "This 
race . . . may be slightly smaller than the other races .... There 
is considerable individual variation, especially as regards the amount 
of black, grey, or reddish brown on the crown and forehead. There 
are female specimens which have a lot of black on the head like the 
males, but specimens occur too without any black markings, or even 
with a reddish crown, which resembles the colour of the back." 
(Schwarz, 1931, pp. 410, 412-413.) 

"The range of this form is only imperfectly known. It clearly 
inhabits the south-eastern coast from Ft. Dauphin in the south to 
at least Masindrano in the north. How far it goes into the interior 
is not known, but it would appear that it is restricted to the coastal 
forests, as L. /. rufus is already found in eastern Betsileo. It also 
remains to be ascertained where the ranges of collaris and julvus 
meet." Specimens are recorded from: Farafangana, about 23 S. ; 
Loholoka, about 21 60' S.; and Fort Dauphin. (Schwarz, 1931, 
p. 412.) 

Ten specimens of "Lemur nigerrimus" were collected by the 
Mission Zoologique Franco-Anglo-Americaine of 1929-31 (Delacour, 
1932, p. 220). 

"At Vondrozo, in June and July, this lemur was common in the 
rain forest, usually traveling in the trees in parties of from four 
to six. . . . 

"The animals were not shy and could be closely approached .... 

"Several stomachs examined contained woody fruits of forest 
trees and one, green vegetable matter that was possibly leaves. 

"At Manomba in October many females were carrying young." 
(Rand. 1935, p. 97.) 


The same author describes the native method of capturing these 
lemurs by means of snares arranged on "a line of poles set up on 
forked sticks across a long, narrow clearing, forming a bridge from 
the trees on one side of the clearing to those on the other." 

[In view of the still existing uncertainty concerning the taxonomic 
status of Prosimia flavijrons J. E. Gray, the exact type locality of 
which is unknown, it seems hardly worth while to give a separate 
account of it here. Schwarz (1931, p. 412) , after provisionally con- 
sidering it a synonym of L. m. collaris, proposes later (1936, p. 24) 
to recognize it as a distinct subspecies of L. macaco and assigns 
to it a range at Maromandia, northwestern Madagascar. On the 
other hand, G. M. Allen (1939, p. 127) regards it as "probably a 
synonym of Lemur macaco collaris" whose range lies at approxi- 
mately the opposite end of Madagascar from Maromandia.] 

Red-fronted Lemur 


Lemur Rujus Audebert, Hist. Nat. Singes et Makis; Makis, p. 12, 1800. (Type 

locality not stated = Madagascar.) 
SYNONYM: Lemur rufifrons Bennett (1833). 
FIGS.: Audebert, 1800, Makis, pi. 2; Milne Edwards and Grandidier, 1890, 

pis. 138, 139, 145. 

This is apparently one of the more common of the Madagascar 

Female (type of rujus) : similar in size to L. mongoz, but differs 
in shorter ears, shorter hairs on tail, and rufous pelage; snout, and a 
line from forehead to crown, black; crown, temples, cheeks, and 
throat dirty white; body yellowish rufous; tail brown at the tip 
(Audebert, 1800, Makis, p. 12) . Male (type of "rufifrons") : back 
grizzly; tail darker; under parts, haunches, and limbs mixed with 
rufous; forehead and cheek-tufts rufous; a nearly complete circle 
of white about the eye; nose, and line through middle of forehead, 
black (Bennett, 1833, p. 106) . The male differs from males of other 
subspecies of L. julvus in having a rusty-red forehead (Schwarz, 
1931, p. 410). This form is very variable in coloration. 

Schwarz (1931, pp. 413-414) describes its occurrence as follows: 

The range of L. f. rujus includes the greater part of Madagascar, all the 
central plateau, and the west coast. ... It would appear that, except the 
north and a fairly narrow strip on the east coast, the whole mainland of 
Madagascar is inhabited by this race. 

As a matter of fact the majority of the individuals found north of the 
Betsiboka River are black-headed and much like the black-headed phase of 
L. /. albijrons; but there can be no doubt that a mixed population is found 
in a considerable part of the north. At present I am not prepared to say 


definitely whether this is due to mutation or, what is more probable, to 
secondary invasion of the northern area by L. /. rujus down the right bank 
of the Betsiboka River. Red-fronted skins have been recorded from north 
of the River Betsiboka from Betsako . . . , Narendry Bay . . . , Anoront- 
sanga . . . ; also . . . from Ambatondrazaka, south of Lake Alaotra. Black- 
headed skins have been recorded from the same general region, but not south 
of the River Betsiboka. In various cases black- and red-fronted skins are 
known from the same localities. 

Additional localities from which Schwarz records specimens are: 
Ankona Forest, E. Betsileo; Fianarantsoa ; Tulear, SW. coast; and 
Morondava, W. coast. 

Lorenz-Liburnau (1898, p. 448) records 16 specimens from Kan- 
dam and 3 from nearby Antema in the Bay of Bombetoka region. 

In the forests in the vicinity of Betsina, west of the Mahavavy 
River, northwestern Madagascar, this lemur is not rare, and four 
specimens were secured (Kaudern, 1915, p. 45) . 

G. M. Allen (1918, p. 515) records "a fine series of six males and 
ten females, all from localities on the upper Siribihina River [inland 
from Morondava] and some thirty miles south of Berevo." 

"This lemur was common about Tabiky [inland from Cape St. 
Vincent] in the low dense brush, the wooded plains, and the gallery 
forest." On one occasion a party of six was seen. (Rand, 1935, 
p. 98.) 

Mayotte Lemur 


Lemur mayottensis Schlegel, Nederl. Tijdschr. Dierk., vol. 3, p. 76, 1866. 

("L'ile de Mayotte," Comoro Isles.) 
FIG.: Schlegel and Pollen, 1868, pi. 2. 

In former years the Mayotte Lemur was apparently common, but 
with the increase of population and cultivation on the island its 
numbers have probably declined. 

Coloration of the head similar to that of L. /. collaris but much 
more pronounced; in old males the snout is deep black, this colora- 
tion extending between the eyes to the forehead and continuing as 
a median stripe to the crown; this color pattern of the head less 
pronounced in younger males and in females; upper parts grayish 
brown, speckled with rufous and yellowish gray; rump with a 
blackish spot; lower parts pale rufous (Schlegel, 1866, pp. 76-77). 
Total length, 39 inches; tail, 21 inches (Schlegel, 1876, p. 308). 

These animals live in bands of 6 to 20 individuals in the virgin 
forests of Mayotte, where they are hunted with dogs. The flesh is 
excellent and tastes like that of young rabbits. A favored food of 
the lemurs is wild dates. A series of 10 specimens is recorded. 
(Schlegel and Pollen, 1868, pp. 5-6.) 


Mongoose Lemur 


[Lemur] Mongoz Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 44, 1766. (Based upon 
the "Mongooz" of Edwards (1758, p. 12); type locality, "Madagascar.") 

SYNONYMS : Lemur nigrijrons, L. albimanus, and L. anjuanensis of fi. Geoffroy- 
Saint-Hilaire (1812) ; Prosimia micromongoz, P. bugi, P. brissonii, and 
P. ocularis of Lesson (1840) ; Lemur cuvieri Fitzinger (1870) ; Propithecus 
brissonianus J. E. Gray (1870). 

FIGS.: Edwards, 1758, pt. 1, pi. 216; Schreber, Saugthiere, vol. 1, pis. 39 B, 42 
("Lemur Simia-Sciurus") , 1774; Audebert, 1800, Makis, pi. l re ; Geoffrey 
and Cuvier, Hist. Nat. Mamm., livr. 2, pi. 11, 1819, livr. 30, pi. 176, 1821; 
Proc. Zool. Soc. London 1871, pi. 16; Milne Edwards and Grandidier, 
1890, pis. 156, 157, 162, 163, 164, 165 (figs. 1, 2); Elliot, 1913, vol. 1, pi. 4, 
lower fig. ("Lemur nigrijrons"). 

In 1915 this lemur was reported as found now and then in the 
forests of northwestern Madagascar, but as apparently nowhere 
common (Kaudern, 1915, p. 43). Concerning its present numerical 
status on certain islands of the Comoro group we have no in- 

Size less than that of a small cat; area about eyes and tip of 
nose black ; area beneath eyes white ; upper parts dark brownish ash 
color; under parts white; tail long (Edwards, 1758, pt. 1, p. 12). 
A white- and a red-cheeked phase, generally sex-linked ; anal region 
almost naked. "In the red-cheeked phase there is not the black 
crown-patch found in the male of L. m. coronatus, whereas in the 
white-cheeked phase the colour of the cheeks is brighter than in the 
female of L. m. coronatus, and a large black or blackish crown- 
patch developed which is absent in that race. The tail is grey or 
blackish in both sexes in mongoz; in coronatus it is reddish in the 
female. As far as my experience goes the males invariably have 
red and the females white cheeks in all specimens from Anjouan 
and Moheli, Comoro Island [s]." (Schwarz, 1931, pp. 414-415.) 

"This race is found on the Comoro Islands, Anjouan and Moheli, 
but not on Mayotte .... On the mainland of Madagascar this 
form is found on the south bank of the Betsiboka River, which it 
ascends up to its head-waters." Specimens are recorded from: 
Anjouan and Moheli Islands; Antema, Bay of Bombetoka, S. bank; 
and Ambatondrazaka, south of Lake Alaotra. (Schwarz, 1931, 
p. 416.) 

Lorenz-Liburnau (1898, pp. 450-451) records 14 specimens (as 
L. albimanus} from Kandani, and 5 from Antema, on the south side 
of the Bay of Bombetoka. 

G. M. Allen (1918, p. 515) records a specimen from Didy, soufh 
of Lake Alaotra. 

Petit (1931, p. 560) records a specimen from Ambongo, north- 
western Madagascar. 


Delacour (1932, p. 219) records 18 specimens of "Lemur nigri- 
frons" collected by the Mission Zoologique Franco-Anglo-Ameri- 
caine of 1929-31. 

Crowned Lemur 


Lemur coronatus J. E. Gray, Ann. Mag. Nat. Hist., ser. 1, vol. 10, no. 65, 
p. 257, 1842. ("Madagascar"; type locality restricted by Schwarz (1931, 
p. 416) to "Bay of Mahajamba, N.W. coast (15 14' S.).") 

SYNONYM: Lemur chrysampyx Schuermans (1848). 

FIGS.: J. E. Gray, Zool. Voy. Sulphur, Mammalia, pi. 4, 1844; Schuermans, 
Mem. Couronnes et Mem, Savants Strangers, Acad. Roy. Belgique, vol. 2, 
pi. facing p. 6, 1848; Milne Edwards and Grandidier, 1890, pis. 158-161, 
165 (figs. 3, 4), 166. 

This subspecies of northern Madagascar apparently still remains 
very common. 

"Ashy above, limbs and beneath pale yellowish; face white; 
orbits gray; cheeks and forehead bright rufous, with a large black 
spot on the crown; tail thick, end blackish" (J. E. Gray, 1842, 
p. 257). Males with red, females with whitish cheeks; anal region 
thickly haired; tail reddish in the female (Schwarz, 1931, pp. 414- 

"This race of L. mongoz is found in northern Madagascar, both 
in the east and west, north of the bays of Bombetoka and Antongil 
respectively." Specimens are recorded from: Vohemar, NE. coast; 
Ampasimbato, Central N. Madagascar; Amber Mountains; and Bay 
of Mahajamba. (Schwarz, 1931, p. 416.) 

Lorenz-Liburnau (1898, p. 449) records one specimen from Betsako 
and four from near-by Ambundube, north of the Bay of Bombetoka, 
under the name of L. mongoz nigrijrons. According, however, to 
Schwarz (1931, p. 416), the animals of this area are coronatus. 

Delacour (1932, p. 219) reports 39 specimens collected by the 
Mission Zoologique Franco-Anglo-Americaine of 1929-31. Judged on 
this basis, it is one of the commonest lemurs of Madagascar. 

"Very common in the dry wooded areas of the northern savanna, 
sometimes in rather low dense brush ; found also in dry forest on the 
slopes of Mt. d'Ambre, up to about 800 meters. It was absent, 
however, from the humid forest on the summit. . . . 

"The animals were very tame and the natives sometimes killed 
them with sticks. . . . 

"Near Vohemar, DuMont and I saw a party of seven .... 

"At Tarakibany ... I saw a party of five." (Rand, 1935, p. 98.) 


Red-bellied Lemur 

LEMUR RUBRIVENTER I. Geoffroy-Saint-Hilaire 

Lemur rubriventer I. Geoffroy-Saint-Hilaire, Compt. Rend. Acad. Sci. [Paris], 

vol. 31, p. 876 (1850), 1851. ("Madagascar.") 
SYNONYMS: Lemur flaviventer I. Geoffroy-Saint-Hilaire (1851); L. rujiventer 

J. E. Gray (1870); Prosimia rufipes J. E. Gray (1871). 
FIGS.: Milne Edwards and Grandidier, 1890, pis. 167-170; J. E. Gray, Proc. 

Zool. Soc. London 1872, pi. 69. 

Even 30 years ago this species was considered rather rare on the 
eastern coast of Madagascar (Kaudern, 1915, p. 47) . 

It may be distinguished from all the other lemurs by the rufous- 
chestnut of its under parts, limbs, and ruff; upper parts speckled 
rufous-brown; tail blackish (I. Geoffroy-Saint-Hilaire, 1851, p. 876). 
Elliot (1913, vol. 1, p. 152) adds the following details from the type 
(a male) in Paris: line from forehead, top of nose, and lips maroon; 
head above mixed dark brown and buff; body above chocolate- 
brown; base of tail maroon, rest blackish; total length, 711 mm.; 
tail, 407 mm. Schlegel's measurements (1876, p. 311) are larger: 
total length, 38 inches; tail, 20 inches. 

Schwarz (1931, p. 417) records specimens from the following 
localities: Vohemar, NE. coast; Bay of Antongil; Betsimisaraka 
country, west of Tamatave; Tamatave, NE. coast; Forest of Ankay, 
NE. of Tananarive; Ambohimitombo and Ivohimanitra, N. Tanala 
country; Vinanitelo, "SW." [=SE.] Betsileo; Manakara River, SE. 
coast; Mojanga, Bay of Bombetoka; and Morondava, W. coast. "If 
all these records are correct the range of L. rubriventer includes the 
greater part of Madagascar, not only the eastern forest region, as 
has hitherto been supposed." 

Delacour (1932, p. 219) records 21 specimens collected by the 
Mission Zoologique Franco-Anglo-Americaine of 1929-31. 

"This diurnal lemur was found in small parties similar to the 
groups of L. julvus. One party contained at least four adults and 
five young. To the west of Andapa [in northeastern Madagascar] 
. . . eight young . . . were taken with the adults." (Rand, 1935, 
p. 98.) 

Ruffed Lemur. Vari (Fr.) 


L[emur] Macaco variegatus Kerr, Anim. Kingdom of Linnaeus, p. 86, 1792. 

(Based upon the "Vari" of Smellie's Buffon (vol. 7, pi. 229, 1791?); type 

locality, "the islands of Madagascar and Johanna, and the neighbouring 

countries of Africa" = Madagascar.) 
SYNONYMS: Lemur ruber 1C. Geoffroy-Saint-Hilaire (1812); Prosimia erythro- 

mela Lesson (1840); Lemur varius I. Geoffroy (1851). 


FIGS.: Buffon, Hist. Nat., vol. 13, pi. 27, 1765; Schreber, Saugthiere, vol. 1, 
pi. 40 B, 1774; Audebert, 1800, Makis, pis. 5, 6; Gervais, Hist. Nat. Mam- 
mif., pt. 1, pi. 10, 1854; Milne Edwards and Grandidier, 1890, pis. 123-129; 
Forbes, 1894, vol. 1, pi. 7; Beddard, 1902, p. 542, fig. 259; Elliot, 1913, 
vol. 1, pi. 5, lower fig. (facing p. 158) ; Kaudern, 1915, pi. 3, fig. 1. 

This lemur is apparently still common in northeastern Mada- 

"Mostly white in the body; all the paws, the fore-head, the 
tail, the insides of the thighs, a large blotch on each shoulder, and a 
long narrow patch on the loin before the upper part of the thigh, 
are black .... The muzzle is long and thick, the ears very short, 
and fringed with long flowing hairs, which join the collar, or hairy 
ruff, on the neck, cheeks, and throat." (Kerr, 1792, pp. 86-87.) 
A color phase, described as "Lemur ruber," has a general rufous 
color; head, hands, tail, and abdomen black; a half-collar of white 
on top of the neck (E. Geoffroy-Saint-Hilaire, 1812, p. 159). 
Total length 44 inches, of which the tail makes up nearly half 
(Schlegel, 1876, p. 302). 

Schwarz (1931, p. 418) records specimens from: Tombato River; 
Sakana, opposite the He Ste. Marie; Alumanitra Forest; Bay of 
Antongil; and Ambatondrazaka, south of Lake Alaotra. He adds: 

"Black and white and red specimens have been collected in the 
same place by J. Audebert in the coast region north of the Bay of 
Antongil. Red specimens are recorded from Maroansetra, Bay of 
Antongil; Malewo and Andranofotsy, N.E. coast, north of the Bay 
of Antongil. . . . 

"From the material at hand it would thus appear that Lemur 
variegatus is restricted to the forests of N.E. Madagascar. Its 
northern limit may be about 13 30' S., its southern range has been 
ascertained as the region of Tamatave at about 18 S. . . . On the 
plateau Ambatondrazaka remains the only place known." 

Coquerel writes (1859, p. 462) that this animal is sacred to the 
inhabitants of Tamatave; they say that it worships the sun and 
prays to it every morning. 

Pollen stated (1868, p. 21) that up to that time it had been 
observed only in the forests of the region between Tintingue, Tama- 
tave, and Antananarivo. It was found there in considerable bands, 
living on fruits. 

Kaudern (1915, p. 43) records specimens from west of Fenerive 
and south of Tamatave. 

G. M. Allen (1918, p. 516) records a specimen from 100 miles west 
of Tamatave. 

The Mission Zoologique Franco-Anglo-Americaine of 1929-31 col- 
lected 8 specimens of variegatus and 9 of "ruber" (Delacour, 1932, 
p. 219). 


"Near Maroantsetra (two days northeast) individuals in the red 
phase, the only phase seen there, were common. The variegated 
lemur is diurnal and arboreal, usually seen in pairs." (Rand, 1935, 
p. 99.) 

Weasel Lemur 


L[epilemur] mustelinus I. Geoffrey, Cat. Method. Mamm. [Mus. Paris], pt. 1, 

Primates, p. 76, 1851. ("Madagascar"; type locality restricted by Schwarz 

(1931, p. 420) to "Tamatave.") 
SYNONYMS: Mixocebus caniceps Peters (1875); Lepidolemur microdon Major 

Fios.: Peters, Monateb. Preuss. Akad. Wiss. Berlin 1874, pi. 1 (facing p. 694), 

1875 (as Mixocebus caniceps) ; Milne Edwards, Grandidier and Filhol, 

1897, pi. 255, pi. 259, fig. 1. 

Only four specimens of this species were collected by the Mission 
Zoologique Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, 
p. 220) . This fact, in connection with the meager information con- 
cerning the animal, indicates that it is one of the less common of 
the Madagascar lemurs. 

General color rufous; throat white; forehead and cheeks gray; 
under parts and inner side of limbs yellowish gray; last third of 
tail brown; rest of tail and lower part of limbs yellowish gray. 
Head and body, about 350 mm.; tail, 250 mm. (I. Geoffrey, 1851, 
p. 76.) 

"The range of L. mustelinus includes the moist east and north- 
east of Madagascar, at least as far south as Betsileo and as far 
north as Vohemar. It does not occur in the north-west, where L. 
ruficaudatus is found." Specimens are recorded from: Vohemar; 
Ankay Forest, NE. of Tananarive; Ampitambe and Antsiraka, 
Betsimisaraka country; Ankona Forest; Upper Masiatra River, E. 
Betsileo; and Vinanitelo, "SW." [ = SE.] Betsileo. (Schwarz, 1931, 
p. 420.) 

G. M. Allen (1918, p. 516) records a specimen from Didy, south 
of Lake Alaotra. 

All the localities of this species are on the eastern slope of the 
island, from Vohemar on the north to Vinanitelo on the south ; these 
localities are littoral or belong to the region of the Hauts-Plateaux. 
The altitudes where it is found vary from less than 100 m. to more 
than 1,000 m. It is probably divisible into subspecies not yet de- 
termiried. All alleged records from the west coast really belong to 
L. ruficaudatus. (Petit, 1933, p. 34.) 


Red-tailed Lemur. Hattock (Madagascar) 


LepilemMr ruficaudatus Grandidier, Rev. Mag. Zool., ser. 2, vol. 19, p. 256, 
1867 - 1 ("La cote sud-ouest de Madagascar"; type locality restricted by 
Elliot (1913, vol. 1, pp. 122) to "Morondava, Madagascar.") 

SYNONYMS: Lepilemur dorsalis Gray (1870); L. pallidicauda Gray (1873); 
Lepidolemur leucopus Major (1894); L. edwardsi Major (1894); L. 
globiceps Major (1894) ; L. grandidieri Major (1894) ; L. mustelinus 
rufescens Lorenz-Liburnau (1898). 

FIGS.: Forbes, 1894, vol. 1, pi. 9 (as L. leucopus}] Milne Edwards, Grandidier 
and Filhol, 1897, pi. 256 (as L. m. var. dorsalis), pi. 257, pi. 258 (as L. m. 
var. leucopus), pi. 259 (as L. dorsalis); Lorenz-Liburnau, 1898, pi. 30 (as 
L. m. rufescens); Kaudern, 1915, pi. 1, fig. 3, pi. 2, fig. 2 (as L. m. 
rufescens) . 

This is apparently a rather common as well as widespread species 
in western and southern Madagascar. No less than 36 specimens 
were collected by the Mission Zoologique Franco-Anglo-Americaine 
of 1929-31 (Delacour, 1932, p. 220) . 

General color ashy rufous; head blackish; hind limbs pale ashy; 
tail reddish; throat fulvous; abdomen whitish. Total length, 560 
mm.; tail, 250 mm. (Grandidier, 1867, p. 256.) 

Schwarz (1931, pp. 420-421) records specimens from the following 
localities: Loko-Be, Nosy Be Island, NW. Madagascar; Betsako, 
north bank, Bay of Bombetoka ; Ambundube, near Betsako ; Antema 
and Kandani, south bank, Bay of Bombetoka; Morondava, W. coast; 
Ambolisatra, SW. coast; and Fort Dauphin, SE. coast. "This species 
has also been recorded by Pollen and van Dam from various points 
on the N.W. coast: Anorontsanga, Jangoa River, Ampasindava, all 
at or near the Bay of Ampasi[n]dava; also from the Bay of Maha- 
jamba. The range, therefore, covers the entire west coast, including 
the north-west and south-east, but not the central plateau. The 
northern and eastern limits remain to be ascertained." (P. 421.) 

Schlegel and Pollen write (1868, p. 12) concerning this species 
(under the name of L. mustelinus) that it is very stupid and more 
slothful than Hapalemur griseus. The natives of the northwest say 
that they sometimes kill it in daytime with sticks and eat its flesh. 
It is nocturnal, and its food consists of buds and leaves of trees as 
well as certain fruits. 

Kaudern (1915, p. 74) records several specimens from Ste. Marie 
de Marovoay on the Betsiboka River, northwestern Madagascar. 

1 Petit (1933, pp. 36-37) recognizes three subspecies, including L. r. dorsalis 
Gray and L. r. leucopus Major. However, he attempts to distinguish dorsalis 
from ruficaudatus in part by "a more russet coloration" and by "a constant 
and more distinct dorsal band," in contradiction to Gray's type description of 
dorsalis (1870, p. 135): "Back grey, yellow-washed; dorsal stripe none." As for 
leucopus, Schwarz states (1931, p. 420) that "the type-specimen is in every 
respect typical ruficaudatus" 



In recognizing dorsalis as a subspecies, Petit states (1933, p. 37) 
that it inhabits the Mahafaly and Antandroy districts, and dis- 
appears abruptly with the very distinct limit of the xerophytic 
vegetation near Bevilana, west of Fort Dauphin. The same author 
(1935, p. 474) remarks on its former presence in the Manampetsa 
Reserve in the southwest, but it was not found there in 1932 or 1933. 

Rand (1935, p. 99) records the "Hattock" from Tabiky and 

Family INDRIIDAE: Sifakas, Indri, Avahis 

The Sifakas and their relatives constitute a family of 3 genera 
and 13 forms. All are endemic to Madagascar. Some of the natives 
have a certain degree of superstitious veneration for the elegantly 
attired Sifakas and the Indri, and yet the animals are not altogether 
free from persecution. In numerical status they vary from common 
to rare. Their limited distribution and their uncertain future render 
all of them suitable subjects for inclusion in the present report. 

Diademed Sifaka. Propitheque a diademe (Fr.) 


Propithecus Diadema Bennett, Proc. Zool. Soc. London 1832, p. 20, 1832. 

("Madagascar"; type locality restricted by Milne Edwards and Grandidier 

(1875, p. 300) to "la cote Est de Madagascar." 1 ) 

SYNONYMS: Macromerus typicus A. Smith (1833); Indris olbus Vinson (1862). 
FIGS.: Gervais, Hist. Nat. Mammif., pt. 1, pi. 8, 1854; Milne Edwards and 

Grandidier, 1875, pi. 1. 

Over a hundred years ago this Sifaka was said to be rare, and 
it is apparently still more so today. The Mission Zoologique Franco- 
Anglo-Americaine of 1929-31 collected only three specimens the 
smallest number reported for any of the Madagascar lemuroids 
(Delacour, 1932, p. 220) . 

Face nearly naked; hairs generally long, silky, waved, erect, and 
glossy; ears rounded, concealed within the fur; a yellowish-white 
band extending across the forehead and below the ears to the throat; 
crown, nape, and hands black; shoulders, sides, and lower back 
mixed black and white; limbs, rump, and tail pale fulvous; throat 
like sides, rest of under parts white. Head and body, 21 inches; 
tail, 17 inches. (Bennett, 1832, pp. 20-21.) 

This subspecies is found only between the Bay of Antongil on the 
north and the Masora River on the south [at about lat. 20 S.], 

i Elliot (1913, vol. 1, pp. 169, 171) attempts the impossible in stating that 
Sambava, northeastern Madagascar, is the type locality of this subspecies as 
well as of P. d. sericeus. For the latter he also attempts to switch Sambava 
to the northwest coast. 


in the narrow bands of forests on the eastern slope of the moun- 
tains (Milne Edwards and Grandidier, 1875, p. 300) . 

"It is stated to be rare" (Bennett, 1832, p. 22). 

G. M. Allen (1918, p. 515) records a specimen from Didy, south 
of Lake Alaotra. 

Schwarz (1931, p. 422) mentions specimens from: Tamatave, 
NE. coast ; Andragoloaka, SE. of Tananarive ; and Mananare, Bay of 

Silky Sifaka. Propitheque soyeux (Fr.) 


Propithecus candidus Grandidier, Compt. Rend. Acad. Sci. [Paris], vol. 72, 
p. 232, 1871. ("Les forets au nord de la baie d'Antongil, sur la cote est 
de Madagascar"; type locality restricted by Schwarz (1931, p. 421) to 
"Sahambavany, N.E. coast" of Madagascar.) 

SYNONYM: Pr&pithecus sericeus Milne-Edwards and Grandidier (1872). 

FIG. : Milne Edwards and Grandidier, 18756, pi. 2. 

This Sifaka has a limited range in the northeast of Madagascar, 
and its numbers are apparently few. 

It is distinguished from P. verreauxi by its entirely white color, 
without black crown or ashy spot on the back (Grandidier, 1871, 
p. 232). Muzzle bare, black, with spots of flesh color; pelage gener- 
ally entirely white, with silky reflections and washed lightly with 
yellow; hairs at base of tail and beneath claws rufous; various 
intergradations between this subspecies and P. d. diadema observed 
(Milne Edwards and Grandidier, 1875a, p. 301). Head and body, 
620 mm.; tail, 400 mm. (Milne-Edwards and Grandidier, 1872, 
p. 274). 

Schwarz (1931, pp. 421-422) records specimens from Sahamba- 
vany, NE. coast, and from Antsompirina and Ansandrizina, NE. 
coast (probably on the peninsula which forms the eastern border 
of the Bay of Antongil). "The range of this race includes the 
north-eastern coast ranges of the island from the region of Bemarivo 
(14 16' 30" S.), which is the northernmost locality recorded, to the 
Bay of Antongil ; I suppose the Tsingambala River, at the northern 
end of the bay, will be found to separate the range of candidus from 
that of diadema." 

The Silky Sifakas inhabit the narrow bands of forests covering 
the eastern slope of the mountains in the northeast, between the 
Lokoy River (13 miles south of Sambava) and the Bemarivo River 
(10 miles north of Sambava). They go ordinarily in smaller bands 
than their congeners; scarcely more than three or four are found 
together. (Milne Edwards and Grandidier, 1875a, p. 302.) 

Elliot (1913, vol. 1, p. 171) is in error in placing the range in 
northwestern, instead of northeastern, Madagascar. 


The Mission Zoologique Franco-Anglo-Americaine of 1929-31 col- 
lected six specimens (Delacour, 1932, p. 220) . 

Milne-Edwards's Sifaka. Propitheque d'Edwards (Fr.) 


Propithecus Edwardsi Grandidier, Compt. Rend. Acad. Sci. [Paris], vol. 72, 
p. 232, 1871. ("Les forets situees dans Pouest de Mananzary, a Mada- 
gascar" ; type locality further delimited by Milne Edwards and Grandidier 
(1875a, p. 303) as "la foret de Manampahy," in southeastern Madagascar.) 

SYNONYM: Propithecus bicolor Gray (1872). 

FIG.: Milne Edwards and Grandidier, 18756, pi. 3. 

No information is at hand concerning the numerical status of 
this subspecies. 

It is entirely black, save for an area of rufous-white on each 
side of the loins; face naked and black; ears well developed and 
covered with long hairs. Head and body, 640 mm.; tail, 460 mm. 
(Grandidier, 1871, p. 232.) Milne Edwards and Grandidier (1875a, 
p. 303) give additional details: black areas lightly washed with 
rufous; a narrow band of reddish black separating the light lumbar 
areas; a light reddish spot at the base of the tail; considerable 
variation in coloration evident, some individuals exhibiting an ap- 
proach to P. d. diadema, and others to P. d. holomelas. 

This Sifaka inhabits the narrow bands of forests that partly 
cover the * eastern slope of the mountains between the Rivers 
Masora (about 20 S.) and Matitanana (about 22 S.) (Milne 
Edwards and Grandidier, 1875a, pp. 304-305) . 

Schwarz (1931, p. 422) is in error in placing the type locality east 
of Masindrano, which would be somewhere in the Indian Ocean. 
He records specimens from: Ampitambe, Betsimisaraka country; 
Ivohimanitra, Tanala; Ambohimotombo, N. Tanala; and Vinani- 
telo, "S.W." [=S.E.] Betsileo. "The specimens from Vinanitelo 
would indicate a considerable extension of the range on the central 

Black Sifaka 


Propithecus holomelas Giinther, Ann. Mag. Nat. Hist., ser. 4, vol. 16, p. 125, 
1875. ("Fienerentova" Fianarantsoa, central Betsileo, Madagascar.) 

Very little information about the Black Sifaka has ever been 

Nearly as large as P. d. edwardsi. "Throat and all the lower 
parts covered with dense fine woolly hair. Male with a small patch 
of ferruginous hairs ... in the middle of the chest . . . ; in the 


female this patch is replaced by two smaller ones ... of a whitish 
colour. All the upper parts deep black, except the back of the root 
of the tail, which is brownish. Abdomen greyish brown. A few 
whitish hairs at the extremity of the tail." Head and body, 23 inches; 
tail, 15-16 inches. (Glinther, 1875a, p. 125.) 

Schwarz (1931, p. 423) records specimens from: Fianarantsoa ; 
S. Betsileo; Nandihizana, N. Betsileo; and "Ambavombe," south 
[ = west?] of Fort Dauphin, S. coast. "The range of this race 
appears to be the inland mountain range in the south-east of 
Madagascar, whereas P. d. edwardsi inhabits the coast range. I have 
little doubt as regards the distinctness of the two races." 

Verreaux's Sifaka. Propitheque de Verreaux (Fr.) 


Propithecus Verreauxi Grandidier, Rev. Mag. Zool., ser. 2, vol. 19, p. 84, 1867. 
("Les contrees arides et sablonneuses ou habitent les Antandroui's, les 
Mahfales, et les Antitenes"; type locality restricted by Schwarz (1931, 
p. 424) to "Tsifanihy, Prov. Antandroy, north of Cape Ste. Marie, S. 
Madagascar." "Mananzari," erroneously stated by Elliot (1913, vol. 1, 
p. 172) to be the type locality, is outside the entire range of the species.) 

FIGS.: Grandidier, Album de Tile de la Reunion, vol. 4, pis. 1, 2, 1867; 
Milne Edwards and Grandidier, 18756, pis. 4, 8. 

This Sifaka has been reported as common in Madagascar from 
Flacourt's time (1661) to the present. 

It is similar to P. d. diadema but with a smaller body, longer 
tail, and more whitish pelage; crown and nape rufous-brown; 
shoulders and sides yellowish white; a gray dorsal patch often 
present; limbs and hands white; face black, almost naked, with 
some white hairs; under parts and inner side of limbs white; tail 
white except at base; throat with a small longitudinal rufous spot. 
Head and body, 490-500 mm.; tail, 450-600 mm. (Grandidier, 
1867a, p. 84.) 

Schwarz (1931, p. 424) records specimens from: Tulear, SW. 
coast; Morondava, W. coast; Bemamanga near Morondava; Anta- 
nosy country; and Ankazoabo, central SW. Madagascar. "Accord- 
ing to A. Milne-Edwards and Grandidier the range of P. v. ver- 
reauxi includes the whole south-western part of Madagascar from 
the Tsidsobon River on the west coast to the region of Ft. Dauphin 
in the south-east. Nothing is known of the exact limits of the range, 
neither in the east, where it should meet P. d. holomelas, nor in 
the west, where no record exists for the region between the Manan- 
bolo and Tsidsobon Rivers, where either this race or P. v. deckeni 
should occur." 

Flacourt (1661, p. 153) reported many of these Sifakas in the 


region west of Fort Dauphin. More than two centuries then elapsed 
before they were rediscovered and named by Grandidier. 

They inhabit the arid coasts in the south and the southwest, from 
Andrahoumbe to the Tsidsibon River. They are always found in 
bands of 10 to 12 individuals. They are diurnal and feed upon young 
shoots of trees and upon fruits. (Grandidier, 1867a, p. 84, and 
1867c, p. 313.) 

In 1866 Grandidier encountered this Sifaka some leagues north 
of the village of the Antandroy king, Tsifanihy, in the vicinity of 
Cape Ste. Marie. Here he found it regarded with a certain venera- 
tion by the natives; they objected to his skinning and dissecting a 
specimen in their village, and they buried the remains ceremoniously 
at some distance from the village. Later he secured a large number 
at various points on the southwestern coast, especially along the 
Morondava River, and at one time he had 15 living individuals. The 
natives capture them by means of snares in the clearings. The ani- 
mals never lived long in captivity. (Milne Edwards and Grandidier, 
1875a, pp. 308-311.) 

"This species was common on the Upper Siribihina River [inland 
from Morondava], and at a locality twenty-five kilometers east of 
Tulear" (G. M. Allen, 1918, p. 515). 

Petit (1931, p. 559) records a specimen from the vicinity of An- 
droka, in the Mahafaly country. 

The Mission Zoologique Franco-Anglo-Americaine of 1929-31 
collected 18 specimens (Delacour, 1932, p. 220) . 

"The brown-capped Propithecus was very common in the wooded 
areas of the southwest. A diurnal, arboreal creature, it was usually 
seen in parties of five to eight. . . . 

"They were usually not at all wild and could be watched at close 
range. . . . Near Tulear ... a group in the trees by the roadside" 
was noticed. 

"Green leaves of trees are probably their staple food. . . . The 
animals were often seen to feed on leaves and the young one I kept 
for a time ate them eagerly." (Rand, 1935, pp. 100-101.) 

This Sifaka has become rare in the Manampetsa Reserve in the 
southwest (Petit, 1935, p. 474) . 

CoquerePs Sifaka. Propitheque de Coquerel (Fr.) 


Propithecus Coquereli A. Milne-Edwards, in Grandidier, Rev. Mag. Zool., 
ser. 2, vol. 19, p. 314, 1867. ("Nord-est" = probably northwestern Mada- 

SYNONYM: Propithecus damonis J. E. Gray (1870). 

FIGS.: Milne Edwards and Grandidier, 18756, pi. 6; Forbes, 1894, vol. 1, 
pi. 11; Kaudern, 1915, pi. 1, fig. 1. 


Up to about 30 years ago, Coquerel's Sifaka was very common 
in parts of northwestern Madagascar (Kaudern, 1915, p. 4). On 
the other hand, not a single specimen seems to have been taken by 
the Mission Zoologique Franco-Anglo-Americaine of 1929-31 (Dela- 
cour, 1932, p. 220). 

General color white; a band of reddish brown extending across 
the chest and onto the arms ; a spot of the same color on the upper 
part of the legs; crown, hands, and inner side of the limbs pure 
white; face covered with very short white hairs; tail slender; fur 
woolly. Head and body of the type (a young individual) , 250 mm. ; 
tail, 220 mm. (A. Milne-Edwards, in Grandidier, 1867c, p. 314.) 
Adults have the head, cheeks, nape, and back yellowish white ; loins 
dark reddish gray; sides and pelvic region dirty white; outer 
side of arms and anterior side of thigh dark chestnut-rufous; tail 
reddish gray (Milne Edwards and Grandidier, 1875a, p. 315) . Total 
length of adults, 3 feet 4 inches (Schlegel, 1876, p. 293) . 

This Sifaka is found only on the northwest coast of Madagascar, 
from the south side of Narinda Bay to the north side of the Bay of 
Bombetoka, between the Rivers Loza and Betsiboka. Numerous 
specimens from this region have been examined. (Milne Edwards 
and Grandidier, 1876, pp. 315-316.) 

Lorenz-Liburnau (1898, p. 454) records five specimens from Bet- 
sako and two from near-by Ambundube, north of the Bay of Bom- 

The animal is very common in the forests on the Ankarafantsika 
Plateau, where it wanders about in bands of 3 to 10 or 12 individuals. 
It was observed in several places between the Betsiboka and Maha- 
jamba Rivers. At Ste. Marie de Marovoay, on the Betsiboka, several 
hundred were seen in the wild, and about 60 specimens were shot. 
(Kaudern, 1915, p. 4, map, p. 5.) 

Additional specimens from Island Nosy Komba and from Am- 
batondrazaka, south of Lake Alaotra, are recorded by Schwarz 
(1931, p. 423). "The specimen from Ambatondrazaka . . . shows 
an eastern extension of the range as far as Lake Alaotra." 

Crowned Sifaka. Propitheque couronne (Fr.) 


P[ropithecus] coronatus "Pollen" A. Milne Edwards, Rev. Scientifique, ser. 2, 
year 1, no. 10, p. 224, 1871. (Type locality not stated; restricted by Elliot 
(1913, vol. 1, p. 174) to "Province of Boeny on the Bay of Bombetok, 

SYNONYM: Propithecus damanus Schlegel (1876). 

FIG.: Milne Edwards and Grandidier, 18756, pi. 7. 

About 30 years ago the Crowned Sifaka was reported as common 
(Kaudern, 1915, p. 6), but apparently none were taken by the 


Mission Zoologique Franco-Anglo-Americaine of 1929-31 (Dela- 
cour, 1932, p. 220) . 

General color white; chest washed with brown; crown black (A. 
Milne Edwards, 1871, p. 224). Similar in size to P. v. verreauxi; 
forehead, head, and cheeks varying from dark blackish brown to 
reddish gray; nape and remaining upper parts white, more or less 
washed with rufous on the limbs and at the base of the tail; a gray 
or brown spot on the nape ; tail and hands white ; under parts vary- 
ing from light rufous to very dark rufous-brown (Milne Edwards 
and Grandidier, 1876, pp. 318-319). Total length, 3 feet 5 inches; 
tail, 21 inches (Schlegel, 1876, p. 294) . 

This Sifaka inhabits the Boeny country, which is comprised be- 
tween the sea on the north, the Betsiboka River on the east, and 
the Manzaray [Mahavavy] River on the west. In the south some 
were killed by Crossley not far from the great forest of Manerinerina, 
where he secured a large number of Decken's Sifaka. (Milne Ed- 
wards and Grandidier, 1876, p. 319.) 

Lorenz-Liburnau (1898, p. 453) records 4 specimens from Antema 
and 21 from Kandani. 

A small band was seen in 1906 on the west side of the Betsiboka 
River opposite Marovoay, and two specimens were collected in 1912 
near the coast between the Bay of Bombetoka and the Mahavavy 
River. It is not rare along this part of the coast, and it is very 
common in the great forests on the Boeny Mountains. It does not 
seem to occur east of the Betsiboka or west of the Mahavavy River. 
(Kaudern, 1915, p. 6, map, p. 5.) 

Decken's Sifaka. Propitheque de Decken (Fr.) 


Propithecus Deckenii Peters, Monatsb. Preuss. Akad. Wiss. Berlin 1870, p. 421, 
1871. ("Kanatzi [=Kanatsy], im 18 s. Br. an der Westkiiste von 
Madagaskar" (Peters, 1869, p. 4).) 

FIGS.: Peters, 1869, pi. 1 (as P. diadema) ; Milne Edwards and Grandidier, 
18756, pi. 5. 

This is still a common animal in western Madagascar. 

The (immature?) type female is described by Peters (1871, p. 
421) as having the hands and head yellowish white like the rest of 
the body; lumbar region and sides washed with gray; face black, 
with a whitish spot on the ridge of the snout; tail as long as, or 
longer than, the head and body. Milne Edwards and Grandidier 
add (1876, pp. 313-314) that adults have a little black diadem in 
front of the white crown; upper chest bright rufous; rest of under 
parts reddish white ; a tawny spot at the base of the white tail. Total 
length 42 inches, of which the tail occupies half (Schlegel, 1876, 
p. 295). 


Crossley secured a number of specimens in the Forest of Manerine- 
rina and in the plains north of Ankavandra [lat. 19 15' S.]. These 
Sifakas inhabit the forests scattered here and there in the midst of 
the great Jurassic plains lying between the Mananbolo and Manza- 
ray [ = Mahavavy] Rivers. The Antimailaka natives consider them 
sacred animals and never kill them. (Milne Edwards and Grandi- 
dier, 1876, pp. 313-314.) 

Decken's Sifaka is very common in the forests south of Lake 
Kinkony (near the lower Mahavavy) , and five specimens were ob- 
tained there. It is said not to occur in the forests between this lake 
and Cape Tanjona. The Mahavavy River appears to form its north- 
eastern limit. (Kaudern, 1915, pp. 6-7, map, p. 5.) 

Beravina, 17 10' S., NW. coast, is one of the localities from 
which Schwarz (1931, p. 424) records specimens. 

No less than 30 specimens were collected by the Mission Zoologi- 
que Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, p. 220) . 

"This Propithecus was common in the country from Namo- 
roka to the Mahavavy Rivers, and a number apparently of the 
same subspecies were seen between the Mahavavy and Betsiboka 
Rivers. . . . 

"This is a common, diurnal animal, found in parties of some- 
times as many as nine individuals. It frequented the heavy gallery 
forest, the lower, dryer forest, and at Soala I found a party in the 
coast mangroves. . . . We found them rather tame here and easily 
approached." (Rand, 1935, p. 99.) 

Major's Sifaka 


Propithecus majori Rothschild, Novit. Zool., vol. 1, p. 666, 1894. ("The 
Antinosy country in south-west Madagascar" ; according to Schwarz (1931, 
p. 424), this is the "country of the emigrated Antanosy, S. Central 

FIG.: Rothschild, 1894, pi. 14. 

Apparently no additional information has been secured concern- 
ing this Sifaka since the original series was collected in 1889 and 
described in 1894. 

"Adult. Head and neck black. Face, snout, and ears naked, 
and of a blackish colour, encircled by a broad band of long white 
hairs, joining under the throat, slightly intermixed with darker 
hairs. Rest of fur, including the tail, white on the upper surface, 
back and upper rump dark brown. The large white patch on and 
between the shoulders much grizzled with brown hairs. Upper sur- 
face of hind limbs to just below the knees blackish brown. Inside 


of hind limbs down to the heel also brown, joining the colour of the 
upper surface, thus forming a continuous dark stripe along the 
legs. Inner and upper surface of arms, thumb, and two following 
fingers deep blackish brown; throat, chest, and greater part of 
abdomen deep brown. Size perceptibly larger than that of Propi- 
thecus verreauxi, with the tail longer. 

"A number of specimens, all perfectly alike in colour, were sent 
to me by Mr. Last from the Antinosy country in south-west Mada- 
gascar. The collector also found Propithecus verreauxi Grandid. 
in the same country, some with the back much darker than others, 
but no specimens in any way intermediate between it and P. majori." 
(Rothschild, 1894, p. 666.) 

"There are four more skins exactly like the type in the Tring 

"The original label only says 'Antinosy' (=Antanosy) country. 
It would appear that this means the mountain range round and east 
of Manansoa (23 3' S., 44 50' E.), where J. T. Last was collecting 
in 1889. The specimens of true verreauxi collected by Last and 
also labelled 'Antinosy country' are probably from the plains farther 
west, and collected on the way to or from Tulear, on the S.W. coast. 
Only the original series of this race is known." (Schwarz, 1931, 
p. 424.) 

Perrier's Sif aka 


Propithecus Perrieri Lavauden, Comptes Rendus Acad. Sci. [Paris], vol. 193, 
no. 1, p. 77, 1931. ("Foret d'Analamera, situee au sud-est de Diego- 
Suarez, dans le nord de Madagascar.") 

This Sifaka is known only from a few specimens collected in the 
Forest of Analamera, which covers an area of 5,000 or 6,000 hectares. 

It bears some resemblance to Propithecus verreauxi coquereli of 
western Madagascar but differs especially in its color. The entire 
pelage is velvety black; ears small and glabrous; eyes brown. Head 
and body, 500 mm.; tail, 450 mm. 

It may seem surprising that this species has remained unknown 
until so recently. The explanation lies in the fact that both the 
animal itself and the forest in which it lives are strictly taboo to the 
local Antakara natives. They give it the name of "Radjako"; this 
was the name of a legendary hero among their ancestors. Few 
Europeans have penetrated the Forest of Analamera. (Lavauden, 
1931, pp. 78-79.) 

G. M. Allen (1939t>, p. 133) regards the type specimen as "prob- 
ably a melanistic individual of P. v. coquereli." 


Inclri; Indris; Endrina. Indri (Fr.) 

INDRI INDRI (J. F. Gmelin) 

[Lemur} Indri J. F. Gmelin, Syst. Nat., vol. 1, pt. 1, p. 42, 1788. (Based upon 

the "Indri" of Sonnerat (Voy. Indes or. et Chine, vol. 2, p. 142, 1782) ; 

type locality, "Madagascar.") 
SYNONYMS: Indris brevicaudatus E. Geoffroy (1796); Indri niger Lacepede 

(1800) ; Indris ater I. Geoffroy (1825) ; Lichanotus mitratus Peters (1871) ; 

Indris variegatus Gray (1872). 
FIGS.: Sonnerat, 1782, vol. 2, pi. 88; Audebert, Hist. Nat. Makis, pi. 1, 1800; 

Milne Edwards and Grandidier, 1875b, pis. 11, 12; Royal Nat. Hist., vol. 1, 

p. 204, fig., 1893-94; Forbes, 1894, pi. 12. 

The distribution of the Indri appears to be subject to considerable 
local variation, but here and there in eastern Madagascar it is still 

This is the largest of the lemurs; it is almost entirely black; fur 
silky and dense; snout, posterior under parts, back of thighs, and 
lower arms grayish; rump white, with woolly hair; tail perceptible 
only to the touch (Sonnerat, 1782, vol. 2, p. 142; J. F. Gmelin, 1788, 
vol. 1, p. 42). It exhibits "a great variety of color pattern as well as 
diversity of hues" (Elliot, 1913, vol. 1, p. 177). Head and body, 
650 mm.; tail, 60-70 mm. (Milne Edwards and Grandidier, 1875a ; 
p. 337). 

Elliot (1913, vol. 1, p. 176) gives the range as "eastern coast of 
Madagascar, in forests on the eastern side of the high mountains 
between the Bay of Antongil on the north and the River Masara 
on the south." 

According to Sonnerat (1782, vol. 2, p. 142), the natives of the 
south capture the animals when young, rear them, and train them 
like dogs for hunting. 

Pollen writes (1868, pp. 20-21) that up to that time the Indri 
was known only from the interior of eastern Madagascar. His 
friend Dr. Vinson reported that while passing through the great 
forest of Alanamasoatrao he was deafened, during two days,' by the 
incessant clamor of apparently numerous but invisible bands of 
these animals. The natives have a superstitious veneration for the 
Indri, and it plays quite a part in their folklore. It feeds upon fruits 
and also preys upon small birds. 

Milne Edwards and Grandidier state (1875a, pp. 340-341) that 
the species lives only on the eastern slope of the great massif be- 
tween the Bay of Antongil and the River Masora. It is essentially 
diurnal and lives in bands, usually of no more than 4 or 5 indi- 
viduals. They refute Sonnerat 's tale of its being trained by the 
natives for hunting. 

"This Lemuroid is probably the best known to travellers in 
Madagascar, at least by ear, as no one can travel along the most 


frequented route in the island, that from Tamatave to Antanana- 
rivo, without often hearing the cries of these animals as he passes 
through the great forest. They are not often seen." (Forbes, 1894, 
vol. 1, p. 109.) 

Schwarz (1931, p. 425) records specimens from the following 
localities: Vohemar, NE. coast; Lalo River and Antsompirina, east 
of the Bay of Antongil ; Sakana, opposite the He Ste. Marie ; Tama- 
tave; Antsihanaka Forest, Lake Alaotra. 

Specimens to the number of 16 were collected by the Mission 
Zoologique Franco-Anglo-Americaine of 1929-31 (Delacour, 1932, 
p. 219). 

Rand writes as follows (1935, pp. 101-102) : 

We encountered the indri only in the heavy forest of the northeast, from 
sea level to 1800 meters, and found some surprising discrepancies in their range. 
About the Bay of Antongil they were common forty kilometers northwest 
of Maroantsetra, but at Maroantsetra, two days northeast, altitude 1000 
meters, where the forest was equally heavy and continuous with that near 
Maroantsetra, none were found. Again, east and north of Andapa there is 
magnificent humid forest but none were heard in it, although west of Andapa, 
perhaps eight kilometers away in similar forest continuous with the former, 
they were common. . . . 

The flesh of these creatures was well flavored but usually so tough and 
hard even when the animals were fat that it was rather unsatisfactory food. 

Many writers have said that this species is sacred to the Malagash. This 
certainly is not true for the Malagash as a whole, for the people of the 
south who had migrated to this part of the island had no objection to 
skinning or eating these creatures, and even the native Betsimisaraka and 
Tsimihity were quite ready to assist us in locating and shooting them, though 
the Tsimihity at Andapa, one day west, would not eat the flesh. 

Eastern Woolly Avahi. Avahis laineux oriental (Fr.) 


[Lemur} laniger J. F. Gmelin, Syst. Nat., vol. 1, pt. 1, p. 44, 1788. ("Mada- 
gascar." Lorenz-Liburnau (1898, p. 452) restricts this subspecies to the 
east coast, and refers to Milne Edwards and Grandidier (1875a, 6), whose 
pi. 9 represents a specimen from the Bay of Antongil. The vicinity of 
this bay may be considered the restricted type locality.) 

SYNONYMS: Lemur brunneus Link (1795); Lemur lanatus Schreber (1800?); 
Indris longicaudatus fi. Geoffrey (1812) ; Semnocebus avahi Lesson 
(1840); Avahis laniger orientalis Lorenz-Liburnau (1898). 

FIGS.: Sonnerat, Voy. Indes or. et Chine, vol. 2, pi. 89, 1782; Schreber, Saug- 
thiere, vol. 5, pi. 42 A, 1800(?); Gervais, Hist. Nat. Mammif., pt. 1, pi. 7, 
1854; Proc. Zool. Soc. London 1866, pi. 15; Milne Edwards and Grandidier, 
18756, pi. 9; Forbes, 1894, pi. 10; Elliot, 1913, vol. 1, col. pi. 7. 

At the present time this appears to be one of the less common 
of the Madagascar lemurs. 

The hair is long and woolly, mouse gray at the base, reddish 
brown in the middle, black at the tips; face broad, covered with 


grayish-brown hairs; nose-pad alone nude; ears concealed, rufous; 
a whitish band across the forehead, bordered anteriorly by a black 
band ; back grayish brown ; a patch over the rump and region about 
the base of the tail white, washed with rufous; under parts and 
inner surface of limbs gray, washed with rufous; tail bright dark 
red, deepest at its extremity (Forbes, 1894, p. 95). Body, 300 mm.; 
tail, 390 mm. (Milne Edwards and Grandidier, 1875a, p. 327). 

"A. I. laniger inhabits the whole forest region of north-eastern, 
eastern, and south-central Madagascar." Specimens are recorded 
from the following localities: Vohemar, NE. coast; Lakato Forest, 
Ankay, NE. of Tananarive; Ambohitra, Kolaby Forest, N. Betsileo; 
Vinanitelo, "S.W." [-S.E.], Betsileo; Fianarantsoa, central Bet- 
sileo. (Schwarz, 1931, p. 426.) 

According to Pollen (1868, p. 21), this species appears to be more 
common than the Indri. It is recorded from the lie Sainte-Marie as 
well as from the mainland of Madagascar. 

The Avahi does not live in bands, but is always found singly or 
in pairs. Its diet is exclusively vegetable. (Milne Edwards and 
Grandidier, 1875a, p. 329; 1875b, map, pi. 122.) 

"The first specimen . . . was brought to Europe by Sonnerat 
. . . in 1781, and nearly half a century elapsed before a second one 
was obtained" (Forbes, 1894, p. 96). 

G. M. Allen (1918, p. 515) records a specimen from the Eastern 

The Mission Zoologique Franco-Anglo-Americaine of 1929-31 
obtained only 9 specimens (Delacour, 1932, p. 219) . 

Rand (1935, p. 102) records several individuals, including a party 
of three, from the vicinity of Vondrozo in the southeast. 

Western Woolly Avahi. Avahis laineux occidental (Fr.) 


Avahis laniger occidentalis Lorenz-Liburnau, Abh. Senckenb. Naturf. Ges., 
vol. 21, p. 452, 1898. ("Ambundube," near Betsako, near Majunga, north- 
western Madagascar.) 

FIG.: Milne Edwards and Grandidier, 18756, pi. 10. 

According to native report, this animal is not uncommon on the 
Ankarafantsika Plateau and on the Bongolava of northwestern 
Madagascar (Kaudern, 1915, p. 2). 

Upper parts gray, with a yellowish-brown shade; woolly hair at 
the base of the tail thin; tail reddish brown, toward the end more 
blackish brown ; hands and feet yellowish brown ; face whitish ; un- 
der parts cream-colored. Body, 330 mm.; tail, 195 mm. (Lorenz- 
Liburnau, 1898, p. 452) . 

Milne Edwards and Grandidier wrote (1875a, p. 329) that this 


western form occurs between Mount d'Ambre and Anorontsangana, 
near the Bay of Passandava. Since then the range has been extended 
considerably to the southward. 

"Collected by van Dam at Kakamba and Ampasidava, N.W. 
coast, by Voeltzkow and [ = at] Ambundube near Betsako, and by 
Kaudern from the Mahajamba River near Ste. Marie de Marovoay. 
The range of this race includes the north-west coast as far south as 
the Bay of Bombetoka; the northern and eastern limits are not 
certain." (Schwarz, 1931, p. 427.) 


The single representative of this family, the remarkable Aye-aye, 
occurs in Madagascar, where it is decidedly rare. 



[Sciurus] madagascariensis J. F. Gmelin, Syst. Nat., vol. 1, pt. 1, p. 152, 
1788. (Based upon the "Aye-aye" of Sonnerat (Voy. Indes or. et 
Chine, vol. 2, p. 137, 1782) ; type locality, "in occidental! parte insulae 

SYNONYM: Lemur psilodactilus Schreber (Saugthiere, vol. 4, pi. 38 D, 1800?); 
Cheiromys madagascariensis var. laniger G. Grandidier (1929). 

FIGS: Sonnerat, Voy. Indes or. et Chine, vol. 2, pi. 86, 1782; Schreber, 
Saugthiere, vol. 4, pi. 38 D, 1800?; Owen, 1863, pis. 14-19; Wolf, 1867, 
pi. 3; Royal Nat. Hist., vol. 1, p. 241, fig, 1893-94; Forbes, 1894, pi. 1; 
Beddard, 1902, p. 548, fig. 263; Lydekker, 1903, frontisp.; Elliot, 1907, 
p. 552, fig. 80; EJliot, 1913, vol. 1, pi. 1; G. Grandidier, Bull. Acad. 
Malgache, n. s., vol. 11, pi. facing p. 101, (1928) 1929. 

The Aye-aye is perhaps the rarest as well as the most interesting 
of all the surviving lemurs of Madagascar. 

The head is short and round; patches of bristles above eyes and 
nose and on cheeks and chin; eyes round, prominent; ears large, 
rounded, naked, black; tail bushy, with hairs 3-4 inches long; 
middle digit attenuated and wirelike. Fur on back, flanks, tail, 
and limbs dark brown, nearly black; long hairs on top of head and 
back of neck tipped with white; face, throat, under parts of body, 
and inner side of limbs yellowish white; feet and digits black; tail 
often with long white hairs throughout. Head and body, 18 inches; 
tail, 18 inches. (J. F. Gmelin, 1788, p. 152, and Forbes, 1894, 
pp. 14-16.) 

"I have not seen any specimen with definite locality. The range 
of this animal appears to include the whole forested portion of 
Madagascar in the east, and apparently also in the north-west." 
(Schwarz, 1931, p. 427.) 

"I am told that the Aye-aye is an object of veneration at Mada- 



gascar, and that if any native touches one, he is sure to die within 
the year; hence the difficulty of obtaining a specimen. I overcame 
this scruple by a reward of 10." (H. Sandwith, in Owen, 1863, 
p. 38.) 

This animal inhabits by preference the bamboo forests of the 
interior. According to the natives, it is very rare; it lives solitarily 

JE, . 

FIG. 18. Aye-aye (Daubentonia madagascariensis) . From photo in Brehm. 

or in pairs; and it is essentially nocturnal. It feeds on the pith of 
bamboos and sugarcane and also on beetles and their larvae. 
(Pollen, 1868, p. 22.) 

"The Aye-aye lives in the dense parts of the great forest that 
runs along the eastern border of the central plateau of the island, 
but only in that part of it which separates the Sihanaka Province 
from that of the Betsimisaraka, and which is about twenty-five 
miles from the east coast, in latitude 17 22' S. or thereabouts. . . . 
From what I have gathered from the natives, it seems to be pretty 
common, its nocturnal habits and the superstitious awe with which 
it is regarded . . . accounting for its apparent rarity .... 

"Occasionally it is brought to Tamatave for sale, where it realizes 
a good sum. Now and then it is accidentally caught in the traps 
which the natives set for Lemurs." (Baron, 1883, pp. 639-640.) 


"Many of the Betsimisaraka still believe that the Haihay is the 
embodiment of their forefathers, and hence will not touch it, much 
less do it an injury. It is said that when one is discovered dead in 
the forest, these people make a tomb for it and bury it with all the 
formality of a funeral." (G. A. Shaw, 1883, p. 45.) 

"It was first discovered by Sonnerat during his travels in Mada- 
gascar in 1780, and by him sent to Paris. The skin remained unique 
in Europe for the best part of a century. ... It was for a long 
period, and is still, very difficult to procure, or to induce the natives 
to capture, specimens." (Forbes, 1894, pp. 16-17.) 

Elliot (1913, vol. 1, p. 2) gives the range as "east coast from 
Bay of Antongil to Mahanoro." 

Kaudern (1915, p. 1) records four specimens from the forests west 
of Fenerive and Tamatave on the east coast. He also mentions 
(p. 2) some questionable reports of the species on the Ankara- 
fantsika Plateau in the northwest. 

G. M. Allen (1918, p. 516) records a specimen from Fenerive on 
the Maningory River. 

"The only aye-aye seen [by the Mission Zoologique Franco- 
Anglo-Americaine of 1929-31] was in the northwest. Throughout 
the rain forest of the east we found few who knew this creature, 
but in the Sambirano it was well known to the natives by name, 
though few had seen it. All of them said it fed on bamboo and was 
very ferocious. . . . Our -single specimen was collected at Ampasa- 
mena, a fishing village on the coast .... This individual ventured 
into the village during the early part of the night and was walking 
about amongst the houses when found by a native, who impaled it 
on a fish spear. It was evidently not common or else not often seen 
as the chief of the village, a gray-haired old man . . . , knew the 
beast by name but had never seen one before." (Rand, 1935, p. 103.) 

In view of the general tolerance and even awe exhibited by the 
natives toward the Aye-aye, its rarity and possibly approaching 
extinction must be attributable to more or less natural causes, as 
yet undetermined. 

Family COLOBIDAE: Leaf-eating Monkeys 

The handsome Colobus Monkeys are externally distinguished 
among African species by the reduction of the thumb, which is 
either very small or altogether absent. A further point of structure 
is in the sacculation of the stomach, a means probably for giving 
greater capacity and a larger absorbing surface to the digestive 
system, for the species are typically leaf-eaters and must in conse- 
quence live upon a type of food requiring bulk and much digestion. 
Two chief types occur, the black-and-white and the red groups. 


These show much local variation, and many names have been ap- 
plied. According to the latest reviser, Schwarz, however, these may 
be regarded as representing but two distinct species, each with 19 
or 20 races, or some 39 in all. They are typically monkeys of the 
great rain forest, from French Guinea south to Angola, and across 
the Congo Basin to the more isolated rain-forest and gallery forest 
of Abyssinia (Ethiopia), Kenya, Tanganyika, and Zanzibar. Over 
a large part of this range both species in one race or another occur 
together, but in some regions only one of them is found, as in Zanzi- 
bar, Kirk's Red Colobus, or in Abyssinia where the black-and- 
white Guereza is alone represented. Both were first made known 
from the West Coast, Sierra Leone. 

G. M. A. 

Black-and-white Colobus; Guereza 

COLOBUS POLYKOMOS (Zimmermann) and races 

Cebus polykomos Zimmermann, Geogr. Geschichte, vol. 2, p. 202, 1780. 

(Sierra Leone.) 

SYNONYMS and list of valid races: Schwarz (1929). 
FIGS.: Elliot, 1913, vol. 3, pis. 3, 18, 19. 

In these handsome black-and-white monkeys, the hair of the 
flanks and hips tends to become elongate, the tail distinctly tufted, 
with progressive increase in amount of white from all-black forms 
(satanas) as in the Cameroons, to those with little and much white. 
The most handsome of the races is perhaps kikuyuensis of Mount 
Kenya or the race caudatus of Mount Kilimanjaro. For the char- 
acters and synonymy of the various races, see Schwarz (1929). 

On account of the long fine hair which forms the prominent 
fringes along the sides, these monkeys are sometimes referred to as 
"shawl monkeys." This quality seems also to have made them 
desirable as fur so that a great many are killed. There is little in- 
formation at hand as to the extent of this trade. Leplae (1925), 
however, states that in the Belgian Congo the fur has a rather high 
commercial value, and the species would be threatened with extinc- 
tion if it were not protected by law. Such protection is given in the 
British colonies but apparently not in the Congo to the extent that 
it should be, although since 1929 it is given partial protection. In 
Kenya Colony the race kikuyuensis occurs and on account of the 
length of its white "shawl" is one of the handsomest of the races. 
Its fur is, or not long ago was, much used by the natives in personal 
decoration. Portions of the black-and-white fur are used as anklets 
(particularly by the young men) or as caps. In the Gabun A. R. 
Maclatchy (in litt., February 5, 1937) found them numerous in 
bands in the mountainous region of Mimongo. They are of sedentary 
habits and affect the high, abrupt mountains. "The vogue which 


their magnificent skin enjoyed lately and even today has been the 
cause of intensive hunting. A furrier of my acquaintance spoke of 
having 30,000 skins in stock, collected from various parts of Africa 
over several years. In view of the animal's restricted habitat, one 
must admit that the protective decree was not unnecessary." These 
monkeys are placed in Schedule B of the London Convention of 1933. 

In general habits the Colobus Monkeys are in the main animals 
of the dense saturate forests; they are not easy to find or shoot 
and will often show considerable adroitness in hiding. Their food 
consists largely of leaves, perhaps also lichens, among the hanging 
festoons of which some of the races live, and probably small forest 
fruits are also taken. Heller has recorded that in the Lado his party 
came upon a troop of Colobus among thorn scrub, to which they had 
come seeking the ripening bean pods, but on being approached they 
made off over the ground to the nearest high forest. Such foraging 
excursions must rather seldom be made in the case of the forms 
which are more strictly high-forest dwellers. Apart from man, their 
enemies are probably limited to leopards and the big crested eagles, 
the food of which consists in part of monkeys. 

While there seems to be little evidence that any of the races is 
at present threatened with extinction, and since the demand for 
their furs seems to have become less, they will no doubt be favored 
by a limited permission to shoot specimens. 

G. M. A. 

Red Colobus 

COLOBUS RADIUS (Kerr) and races 

Simia (Cercopithecus) badius Kerr, Anim. Kingdom of Linnaeus, p. 74, 1792. 

(Sierra Leone, based on the Bay Monkey of Pennant.) 
SYNONYMS and list of valid races: See Schwarz, E., Zeitschr. f. Saugetierkunde, 

vol. 3, pp. 92-97, June, 1928. 
FIGS.: Elliot, 1913, vol. 3, pis. 5, 6, 14-16 (animal and skulls). 

The Red Colobus Monkeys include no less than 20 recognized 
races and differ in color from the black-and-white group, in having 
the fur more or less black and red in varying pattern. Since the 
fur is not as modified in long fringing patches along the sides, it is 
not in special demand. Nevertheless one or two of the races are 
rare or localized and may require special protection for their con- 
tinued safety. 

G. M. A. 

Gordons 9 Red Colobus 


Piliocolobus gordonorum, Matschie, Sitzb. Ges. Naturf. Freunde Berlin, 1900, 
p. 186. (Uzungwe Mountains, Uhehe, Tanganyika Territory.) 


This rather strikingly colored subspecies is known only from a 
circumscribed area in the Uzungwe Mountains to the northeast of 
Lake Tanganyika. The name was based on a single imperfect skin 
found in a native hut and two other skins secured by the brothers 
von Gordon, for whom it is named. In 1923 Kershaw recorded an- 
other specimen secured in the same region by Arthur Loveridge, 
who on a second visit obtained four others for the Museum of Com- 
parative Zoology. 

The top of the head is deep ferruginous, the back shining black; 
forelimbs black, hind limbs mixed black and silvery, the tail mixed 
black and ochraceous; lower surfaces white. 

On the somewhat isolated Uzungwe Mountain range Mr. Loveridge 
found this monkey but once, when at an altitude of some 5,000 feet 
he came upon a troop just at dusk. They live in high forest and are 
with difficulty obtained. Their nearest relative is perhaps Kirk's 
Red Colobus of Zanzibar. Their chief danger is perhaps from native 
hunters, but also possibly in future encroachments upon the small 
area of forest to which they are confined. 

G. M. A. 

Kirk's Red Colobus 


Colobus kirkii Gray, Proc. Zool. Soc. London 1868, p. 180, May, 1868. ("Zan- 
FIGS.: Gray, op. cit., pi. 15; Elliot, 1913, vol. 3, pi. 16 (skull). 

Restricted to the Island of Zanzibar, this form is in danger only 
to the extent that future settlement and cultivation may reduce its 
area of habitat. 

Forehead and sides of head yellowish white, the long hairs extend- 
ing beyond the sides of the head; crown, lower part of neck, back 
from shoulders, reddish brown; shoulders, outer side of arms, hands 
and feet black; throat and under parts grayish white; tail dull 
reddish brown. 

The chief interest of this monkey, apart from its island habitat 
to which no member of the black-and-white group extends, is that 
in the skull the median frontal suture remains open into adult life, 
a rare condition sometimes found also in man. 

This monkey was first secured and sent to Europe by Sir John 
Kirk in 1868. He regarded it as rare at that time but in 1884 wrote 
that it was still to be found in many of the wooded districts of the 
island, although "so rare as not to be procurable, even when I sent 
the hunters over the island. I have a report that it exists still in 
one spot, which they could not reach. ... It looks as if the animal 
will be lost to science. This is due to the destruction of forest and 
jungle over the island." Two years later Sir Harry Johnston wrote 


that it had "disappeared from nearly every part of the island of 
Zanzibar, but a rumor prevailed that -it still lingered on a clump of 
forest as yet unvisited by hunters." On sending his hunters thither, 
they returned after a week's absence, bringing 12 dead monkeys, 
with the report that they had killed every one, so that, as Sir Harry 
supposed, this animal too had gone to "the limbo of species ex- 
tinguished by the act of man." Nevertheless these evidently were 
not the last, and even to this day a few still remain on the island, 
but of their number and present status no information is at hand, 
beyond the fact that Arthur Loveridge procured a pair there in 1923. 

G. M. A. 

Family PONGIDAE: Anthropoid Apes 

The two forms of Gorillas (genus Gorilla) and the four forms of 
Chimpanzees (genus Pan) are found in central Africa. The third 
genus of the family, the Orang-utan (Pongo) , is represented by one 
species, occurring in Borneo and in Sumatra. As man's nearest 
living relatives, these apes have an exceptional interest for us, and 
their generally waning numbers call for a discussion of each form 
in this volume. 

Orang-utan. "Mias" (Borneo) ; "Mawas" (Sumatra) 

Simia pygmaeus Hoppius, Amoenit. Acad., 1763, p. 68. (Locality unknown.) 

SYNONYMS: Simla satyrus Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 34, 1766 (not 

of the 10th ed.) ; Pongo wurmbi Tiedemann, Zool., p. 329, 1808 (Borneo) ; 

Simia morio Owen, Proc. Zool. Soc. London 1836, p. 92, 1837 (Borneo) ; 

Simla abelii Clarke, Asiatick Researches, vol. 16, p. 489, pis. 1, 3, 4. 5, 

1825 (Sumatra). For extensive synonymy, including names given by 

Selenka and others, see Elliot (1913, vol. 3, pp. 192-195). 

FIGS.: Elliot, 1913, vol. 3, pis. 5 (photos of animal), 23-28 (skulls); Carpenter 

and Coolidge, 1938, fig. opp. p. 18; Yerkes and Yerkes, 1929, figs. 43-66. 

It is at present believed that the Bornean and Sumatran Orangs 
are not separable even subspecifically ; at least they represent the 
same species and are not found living elsewhere. No doubt they in- 
habited the Asiatic mainland at no very distant time, but with the 
separation of Borneo and Sumatra from the Malay Peninsula the 
populations of these islands were cut off and have survived to the 
present. On the mainland, remains of anthropoids resembling the 
Orang are known from the Siwalik Hills of India, but there seems 
to be no evidence of their survival to the historic period. 

The adult Orang-utan is a large shaggy animal, of dark rufous 
color. The profile of the skull is much more sloping than in the 
African anthropoids, the skull showing very little of the brow ridges 


so prominent in the latter. The arms are very long, reaching to 
the ankles when the animal is erect; foot long and narrow, the 
great toe very short. Tail absent. Prominent cheek callosities some- 
times present in adult males. Wallace, who measured 17 freshly 
killed Orangs, states that adult males "only varied from 4 feet 1 
inch to 4 feet 2 inches in height, measured fairly to the heel, so as 
to give the height of the animal if it stood perfectly erect; the extent 
of the outstretched arms from 7 feet 2 inches to 7 feet 8 inches." The 
total length of a Bornean skull is said to be 246 mm. (Elliot) , for 
the largest of many. Selenka gives series of measurements. For an 
excellent account of the history, characters, psychology, see Yerkes 
and Yerkes (1929). 

The Orang is a much more lethargic animal than the African 
anthropoids, moving leisurely through the forest, seeking various 
fruits, especially those of the durian, of which it is extremely 
fond. Leaves and bark of certain trees are also eaten. It is some- 
what social and may be found singly or in pairs or in small groups. 
Banks (1931) writes: 

In a wild state and unmolested, Mias exhibit little more than a benevolent 
curiosity towards men and the extremely child-like and almost pathetic 
expressions that can be assumed in captivity point to the Mias as an extremely 
peaceful and gentle animal when left to himself, always remembering of 
course that both temper and strength are there in reserve for use when 
aroused. . . . [They] make a kind of platform of sticks on which they sleep 
at night and even during the day but I have never seen captive ones make 
any sort of roof or make use of leaves to keep the rain off, as is sometimes 
alleged. Nests are of two kinds, either a flat platform or more usually a 
deep triangular shaped affair in the upright fork of a tree. ... I counted 
eleven such nests still with green leaves all close together near a "Kayu 
Ara" fruit tree where a pair were feeding. . . . 

The distribution of the Mias in Sarawak is peculiar in its relations to the 
rest of Borneo; it occurs in parts of N. Borneo . . . and it is common in 
W. Borneo, the Landak River and right up the Kapuas River. Now the 
Mias is very sensibly fond of neither cold nor rain, in fact the damp is his 
worst enemy and for this among other reasons the occurrence of Mias at 3000 
ft. is very exceptional nor is he as common in the immediate lower vicinity 
of mountains as he is at the foot. For some 70 miles the Kalinkang Moun- 
tains run N.E. and S.W. forming a watershed between that part of the Kapuas 
River running S.W. and numerous short Sarawak rivers running West into 
the sea and it is obvious that these mountains form an obstacle to the 
movements of Mias which are common on the Kapuas and curiously on 
the Sarawak side. The explanation lies I think in a gap in the Kalinkang 
Mountains which towards Lobok Antu slope away almost to sea level, 
eventually to rise on the other side of the Batang Lupar Mts. and stretch 
away unbroken northwards into central Borneo. It is therefore more or less 
true that the Mias is confined to a range bounded on the N.E. by the Rejang 
River, on the west by the Sadong River: the Orang Utan has flowed through 
from Dutch Borneo and filled up suitable and available places. 

In upper Sarawak the Orang was formerly reported, but these 
reports are doubtful, although Everett records two imperfect skulls 



found in a crevice of the limestone hills at Paku, but at present 
it is absent from all that region adjacent to the watershed of the 
Landak River. According to Wallace, it has a wide distribution 

FIG. 19. Orang-utan (Pongo pygmaeus). From photo of specimen in 
Philadelphia Zoo. 

in the low country, inhabiting many districts not only on the south- 
west and southeast, but also on the northeast and northwest coasts, 
but of its more exact local distribution at the present time little 
information is at hand. 

In Sumatra, the only place outside of Borneo where the Orang 
is native, it is now confined to the former state of Atjeh, which 


comprises practically the northwestern quarter of the island. In 
their recent report on the animal here, Carpenter and Coolidge 
(1938) indicate that the regions of Lami, Tapa Toean, and Ba- 
kongan are the places on the west coast of Atjeh where Orang-utans 
live in the largest numbers. They occur over the greater part of 
Atjeh except the high central districts above 1,500 meters, the cul- 
tivated and thickly populated districts on the east coast, the grass- 
lands of the north, the rough mountains north of Lamno, and culti- 
vated sections of the west coast, especially around Meulaboh. 
Although an estimate of the numbers and normal density of popu- 
lation is largely guesswork, it seems likely that the centers of abun- 
dance are "around the Simpang Kanan and Peureulak Rivers on 
the East Coast, and along the West Coast in suitable forests from 
Lami to Singkel." They show a marked preference for lowlands 
but may range up to 1,500 meters, though with a sharp decrease 
above 700 or 800 meters. There is some evidence of local move- 
ments following the seasonal ripening of certain fruits on which they 
feed. In summary, these authors state that the Orang is found in 
"an estimated fifty per cent of the primary forests" in Atjeh; "as 
large clearings are made in* the rather level lowlands, these apes 
are being destroyed or forced into the hills and mountains where 
it is questionable whether or not conditions, including food supply, 
are sufficiently suitable for the maintenance of the present popula- 
tion level. However, large areas of Atjeh, because of its rugged 
topography and inaccessibility, will remain naturally protected for 
a long time as an orang-utan habitat. Europeans and not natives 
threaten the orang-utan population, the most serious inroads being 
made by commercial developments in the areas suitable for orang- 
utan habitats. Numbers of these apes are being shot annually under 
the supposition that they attack human beings, and it is feared 
that government records do not accurately record all animals cap- 
tured or killed." 

For the better preservation of these interesting apes, the authors 
recommend an extension of the present Alas National Park or Loser 
Reservation and the development of a smaller reserve especially for 
Orangs south of Meulaboh, together with the planting of food trees, 
especially the durian. Further, the desirability of additional reserves 
and special measures is advocated, particularly that "the killing 
and capture of this animal for trading or exhibition purposes be 
completely stopped and that its use for accredited but limited scien- 
tific purposes alone be permitted." 

G. M. A. 


Coast Gorilla. Gorille (Fr.). Gorilla (Ger.) 


Troglodytes'] gorilla Savage and Wyman, Proc. Boston Soc. Nat. Hist., vol. 2, 
p. 245, 1847. ("Empongwe, near the river Gaboon, Africa"; about lat. 
20' N., long. 9 30' E.) 

FIGS.: Gervais, Hist. Nat. Mammiferes, pt. 1, pis. facing pp. 26, 28, 1854; 
Du Chaillu, 1861, frontisp.; Forbes, 1894, vol. 2, pi. 28; Proc. Zool. Soc. 
London 1896, p. 505, fig.; Elliot, 1913, vol. 3, frontisp.; Cunningham, 
1921, pp. 119-124, figs.; Barns, 1923, figs. 45, 46, 50, 51; Yerkes and Yerkes, 
1929, numerous figs.; Coolidge, 1936, pi. 12; Raven, 1936a, p. 316, fig.; 
Fauna [Philadelphia], vol. 1, no. 1, pp. 8-9, figs., 1939. 

The Coast Gorilla is generally considered to be diminishing in 
numbers but not to be in danger of extermination. All gorillas are 
given full protection under Schedule A of the London Convention 
of 1933. 

"This animal ... is much larger and more ferocious than the 
Chimpanzee. Its height is above five feet; but it is remarkable for 
the disproportionate breadth of the shoulders, which is double that 
of the Chimpanzee. The hair is coarse, and black, except in old 
individuals, when it becomes gray. The head is longer than that of 
an ordinary man by two inches, and is remarkable for having a crest 
of coarse hair over the sagittal suture, which meets at right angles 
a second, extending over the upper part of the occiput, from one 
ear to the other. The fore-arm is much shorter than the arm, the 
hand is remarkable for its great size, and the thumbs larger than 
the fingers." (Savage and Wyman, 1847, pp. 245-246.) 

"Face and chest bare, black; . . . arms and belly black; back 
and outside of thighs gray grading into black towards ankles and 
on feet; hands black; no beard; top of head black, nape mixed 
black and red." Height, 5 feet 10 inches. (Elliot, 1913, vol. 3, 
pp. 213-214.) 

The numerous described forms of Gorillas are reduced by Cool- 
idge (1929, p. 348) to two subspecies the present one and the 
Mountain Gorilla (G. g. beringei). 

"For the Coast Gorilla, the westernmost boundary approximates 
the Cross River in the southern provinces of Nigeria. The most 
westerly point actually recorded is Ikom, 8 40' east and 6 north. 
The northernmost point is close to Basho, 9 25' east, 6 7' north. 
On the east we have reports from several places such as Wesso and 
Nola on the Sanaga [error for Sanga] River. The Sanaga River, 
about 16 15', seems to mark the eastern boundary of the range 
of the Coast Gorilla. On the southeast the line follows the border 
of the forest which reaches its southernmost limit at Mayombe on 
the edge of the Belgian Congo, 5 south, 13' [ = 13] east. Along 
the Atlantic coast in most places the forest begins a little way inland. 


Gorillas have been reported actually on the coast, but generally they 
are found not closer than thirty miles from the sea. They seem 
especially plentiful along the Gaboon, Ogowe, Camp, and Sanaga 
Rivers." (Coolidge, 1929, p. 363.) More recently the range has been 
found to extend somewhat eastward of the Sanga River in French 
Equatorial Africa (Coolidge, 1936, p. 493, maps 1-2). The total 
range of the Coast Gorilla seems to be strictly confined to the west- 
ern portion of the Lower Guinea Forest District of Chapin (1932, 
p. 90) and of Bowen (1933, pp. 256, 258). 

Gabun. "They live in herds, the females exceeding the males in 
number. . . . They are exceedingly ferocious, and objects of terror 
to the natives, who seldom encounter them except on the defensive. 
The killing of an Engeena is considered an act of great skill and 
courage, and brings to the victor signal honor. . . . 

"Their flesh, when obtained, is eaten by the natives, as well as 
that of the Chimpanzee." (Savage and Wyman, 1847, p. 246.) 

Du Chaillu's classical account of the Gorilla (1861, pp. 388- 
404) is based upon his experiences in Gabun. Among other things, 
he says (pp. 399-400) : "The negroes never attack them with other 
weapons than guns; and in those parts of the far interior where no 
European guns had yet reached, as among the Apingi, this great 
beast roamed unmolested, the monarch of the forest." 

"The Fernan Vaz District ... is considered the best region 
for both the gorilla and the chimpanzee." One "family party of 
five or more gorillas" was encountered, and on another occasion 
"a large family" was reported. The animal will occasionally ad- 
vance to attack when not wounded. (Aschemeier, 1921, pp. 90-92.) 

"The majority of the Gaboon skulls have come out from the 
region of the Ogowe River" (Coolidge, 1929, p. 303). 

A. R. Maclatchy (in litt., February 5, 1937) gives the following 
report for Gabun: "The decree of 1929 classed the gorilla among 
the protected species. Its great vulnerability makes it an adversary 
much less dangerous than the buffalo and the elephant. It rarely 
pushes its attack to the limit. Sometimes it visits the native plan- 
tations by night. Its food consists of various plants. I do not see 
the reasons for a protection as strict as that which the gorilla 
enjoys. It is little hunted, except in legitimate defense, by the 
natives, who have a superstitious terror of it, and by a few hardy 
European sportsmen. It scarcely seems to be threatened with de- 
struction. It really abounds, and is protected by its habitat and 
by its natural shyness. More specimens could be allowed on hunting 
permits without the risk of diminishing its numbers." 

Cameroons. In the southeastern corner of Cameroons, and per- 
haps also across the boundary in French Equatorial Africa, the 
Gorilla appears quite common, and almost entirely inoffensive as 


to attacks on man. The forest is so dense, and the animal so shy, 
that it is extremely difficult for the hunter to get sight of it. Thus 
the Gorilla seems fairly well protected. Bands numbering up to more 
than 20 individuals are reported. (Ramecourt, 1936, pp. 217-247.) 
Raven (1936a, 19366) gives a most interesting and detailed 
account of hunting Gorillas during more than a year spent at va- 
rious places in southern Cameroons. The natives here are very keen 
to eat Gorilla meat, being generally faced with a deficiency of 
meat in their diet. A missionary reported many of the animals at 
Djaposten, in southeastern Cameroons, where "in one morning's 
walk of perhaps two hours he had counted more than 100 gorilla 
beds." Although Raven himself found the animals quite common in 
this region, it was extremely difficult to obtain a good view of them 
in the dense forest, and only three adults were collected during his 
entire sojourn, despite assiduous hunting. Raven writes further 
(19366, pp. 529-530) : 

For centuries past the gorillas and natives have been competitors. As the 
native populuation increased, new villages would be formed and more 
clearings made. Then epidemics would occur, killing off great numbers of 
natives, and their gardens would be neglected to run into second growth. 
The gorillas, with a constitution so nearly like that of man that they can 
find more food in human plantations than in the virgin forest, would move 
into these deserted clearings. There with an abundance of food they throve 
and congregated, to such an extent eventually that if only a few natives 
remained they were actually driven out because of their inability to protect 
their crops against the gorillas. But with the advent of the white men's 
government, with the distribution of firearms among the natives, preventive 
medicine and the treatment for epidemic and infective diseases, man has the 
upper hand at present in this age-long struggle. 

"Mr. Raven had opportunity to witness the unfortunate effect, 
so far as the protection of the gorilla was concerned, of the demand 
for gorilla skulls on the part of scientists, to such a degree that 
white men as well as natives had in the past often done a profitable 
business in killing the animals and selling their skulls. The result 
had been a rapid decrease in the gorilla population, so that Mr. 
Raven, although by his record known to be a hunter and collector 
of the first rank, was compelled to hunt week after week in a des- 
perate effort to come up with the nervous survivors of the race in 
this district. . . . 

"Mr. Raven's experience leads him to believe that ... the gorilla 
is being rapidly exterminated in many localities." (Gregory, in 
Raven, 19366, p. 540.) 

It is doubtful if the protective laws have stopped the killing of 
Gorillas by natives to any extent. Most of the museum specimens of 
skulls, etc., are from native-killed animals and have been turned 


in to traders. Natives capture them by spearing and by snaring. 
(H. C. Raven, oral communication, March 17, 1937.) 

The Gorilla is utilized for experimental purposes in the study of 
human diseases (Ministry of Colonies, Paris, in litt., November 7, 

Gorillas are decreasing but not disappearing. The cause of 
depletion is native hunting for food. (Inspection of Waters and 
Forests, Yaounde, in litt., January 12, 1937.) 

The number in French Cameroons is estimated at some thousands. 
They are partially protected by law, one head being allowed on a 
full license. (Paris Agency, in litt., November, 1936.) 

Nigeria. Coolidge (1929, p. 303) refers to the range as including 
"the Western Cameroons [part of Nigeria], which is a comparatively 
limited section centering around Mamfe or Dakbe and extending 
west as far as the Cross River. A great many skulls come from this 

Hay wood (1932, p. 32) reports the species from the borders of 
Ogoja and Cameroons Provinces. 

In British Cameroons, Sanderson (1935, p. 26) reports Gorillas 
from the mountains of Assumbo, about the headwaters of the Cross 
River. They "are numerous in the Mountain Moss Forest belt, 
where the natives record their movements minutely." 

"In Nigeria where a few exist the natives take an annual toll and 
I do not think there are many" (C. W. Hobley, in litt., August 18, 

French Equatorial Africa. The Gorilla does some damage in the 
banana plantations, but it is not important. It seldom attacks man 
without provocation. There is no reason why it should not be abso- 
lutely protected. Its northern and western limits are unknown; its 
southern seems to be the Congo. (Lavauden, 1933, p. 30.) 

"Four complete specimens of the Coast Gorilla were procured 
by the Vanderbilt Expedition of the Academy of Naturaf Sciences 
of Philadelphia in the winter of 1934." Three "were killed by natives 
in the neighborhood of Aboghi, forty miles southwest of Nola near 
the west bank of the Sanga River." The fourth was secured "near 
Barundu, about 15 miles east of the Sanga River and 22 miles north- 
east of Nola." (Coolidge, 1936, p. 479.) 

Green (in Coolidge, 1936, pp. 491-492) reports Gorillas as abun- 
dant in the region of Aboghi. "The old males appear to be somewhat 
solitary, but small bands of four to ten were noted from tracking." 

"The expedition reported that gorillas were frequently killed on 
the left bank [of the Sanga]. . . . 

"The field notes of Mr. Rehn and Mr. Green give us the impression 
that gorillas were plentiful in the region from which these specimens 


came, and that they are frequently hunted by the natives." (Cool- 
idge, 1936, pp. 493, 499.) 

According to all accounts, Gorillas still occur in fair numbers in 
the Sanga River region, though not so commonly as 25 years ago.~A 
local French doctor, in the course of two years, had treated nine 
natives for Gorilla attack, one of the cases being fatal. (J. A. G. 
Rehn, oral communication, March 22, 1939.) 

Belgian Congo. Schouteden (19306, pp. 298-299) presents evi- 
dence of the rare occurrence of Gorillas in the Mayumbe forest, 
north of the lower reaches of the Congo River. Later (19366, 
pp. 15-16) he records a skull from the Haut Mayumbe. Here the 
animal had seemed to have disappeared, or to occur only occasion- 
ally, coming perhaps from Gabun. But it appears to occur still in 
certain parts of Mayumbe, thanks, perhaps, to the protection it has 
enjoyed for some years. 

Use in research. Yerkes and Yerkes point out (1929, p. 590) 
that the Gorilla and other anthropoid apes "must inevitably become 
the preferred substitutes for human subjects in investigations which 
may not be carried on with the latter and which have as objectives 
the extension of knowledge and control of human life." They also 
stress (p. 589) the greater availability and controllability of these 
animals for use in the investigation of various problems in genetics, 
physiology, neuro- and psychopathology, psychology, sociology, 
pedagogy, and experimental education. 

Survival Yerkes and Yerkes (1929, p. 396) quote Keith (1896) 
as follows: "From accounts furnished by travellers and hunters, one 
infers that the total population [of the species as a whole] males, 
females, and young is well under 10,000." Eventually (1914) 
Keith raised this estimate to 20,000 to 30,000 individuals, but 
Yerkes and Yerkes remark (p. 397) that it is difficult to decide how 
seriously this estimate should be taken. They also say (p. 397) : 

"Concerning abundance or frequency little is known. Both early 
and late in the last century the relative rarity of the gorilla sug- 
gested to investigators its disappearance and probable extinction. 
From limited distribution, difficulty of negro hunters in procuring 
skins of adults, and the small number of captive specimens sent to 
Europe, Deniker (1891, pp. 369-370) infers that the process of 
extinction is under way. . . . 

"For nearly a century it has been known that the gorilla is the 
rarest of the manlike apes." 

H. C. Raven (oral communication, March 17, 1937) estimates 
the total number of all Gorillas now living at more than 1,000 and 
at less than 10,000. 


Mountain Gorilla. Gorille ties montagnes (Fr.) 


Gorilla beringeri [misprint for beringei] Matschie, Sitz.-Ber. Ges. Naturf. 
Freunde Berlin 1903, no. 6, p. 257, 1903. ("Auf der Spitze des Vulkans 
Kirunga ya Sabinyo in einer Hohe von 3000 m," German East Africa; 
i. e., Mount Sabinio or Sebyinyo, at the boundary point of Ruanda, 
Uganda, and the Belgian Congo. Not at the summit, but on the south 
or southeast flank, at about 2800 m. (Derscheid, 1928, p. 150).) 

FIGS.: Lonnberg, 1917, pi. 1; Barns, 1922, frontisp. and pis. facing pp. xvi, 
83, 86; Barns, 1923, figs. 43, 44, 52; Akeley, 1923a, pp. 428, 438, 440, 
444, figs.; Akeley, 19236, frontisp. and pis. facing pp. 190, 206, 222, 230; 
Yerkes and Yerkes, 1929, numerous figs.; Coolidge, 1929, pi. 1, and 1930, 
pp. 626-627, figs. 454, 454b; Raven, 1931, cover and p. 241, fig.; Bingham, 
1932, pis. 18, 19, 22; Jour. Soc. Preservation Fauna Empire, n. s., pt. 18, 
frontisp., 1933. 

The Mountain Gorilla is now well protected in its range centering 
in the Pare National Albert in the eastern Belgian Congo, and its 
chances for survival appear to be excellent. 

Face, ears, breast, back, hands, and feet naked; breast brownish, 
like worn leather; back somewhat lighter; face, ears, and naked 
parts of the limbs black; hair black, long, and thick, and forming a 
pronounced beard on cheeks and chin. Height 1.5 m.; weight 100 kg. 
(Matschie, 1903a, p. 254.) "The external characters that distinguish 
the Mountain from the Coast Gorilla are, besides a longer palate 
and a generally narrower skull, the thicker pelage, shorter arms 
and longer legs, large amount of black hair, and fleshy callosity on 
the crest" (Coolidge, 1929, p. 375). "The large patch of silver-gray 
fur covering the back of the adult male gorilla is the most remark- 
able part of his coloration; the female is entirely black, and very 
much smaller than her mate" (Barns, 1923, p. 130) . 

Coolidge (1929, p. 363) says of its range: 

The Mountain Gorilla is found in a comparatively narrow strip of the 
eastern Congo. Its principal habitat is the mountain forest as distinguished 
from the lowland forest of the Belgian Congo. Its northern limit is Mulu, 
10' south, 29 10' east (Absil and Chapin). We find it as far west as 
Walikale, 1 20' south, 28 1' east, where it strays a little into the lowland 
forest. The eastern limit seems to be close to Kigezi in Uganda, 1 15' south, 
29 45' east. The southern limit is Baraka on Lake Tanganyika, 4 19' south, 
29 2' east. In this entire region the gorillas that are most known and accessible 
are the troops that inhabit the volcano regions where Akeley died while 
studying them. Whether they are entirely isolated from contact with outside 
gorillas at the present time is doubtful and has not yet been established. 
In the mountains back of Baraka, Boko, Uvira, and Katana large troops 
have been recently found in the upland forests. 

"I have examined . . . the sources of evidence for the existence 
of gorillas in the intervening area between (longitude 17 east) the 
eastern limit of the known range of the Coast Gorillas and (longi- 


tude 28 east) the western limit of the known range of the Mountain 
Gorillas. With a single exception, I attach no great importance to 
this evidence. The exception refers to the four skulls from Bondo 
on the Uelle River collected by Lemarinel in 1908. These furnish 
us with definite proof for the existence of gorillas in the Djabbir 
region as late as 1908. Except for these skulls no other tangible 
evidence of gorillas in a forest belt of 650 [ = about 750] miles has 
turned up." (Coolidge, 1936, p. 500.) Coolidge considers (p. 497) 
that the affinities of the Bondo skulls are with the Coastal Gorilla, 
but G. M. Allen (1939, p. 177) refers G. uellensis Schouteden, which 
was based upon these skulls, to the synonymy of beringei. 

The Mountain Gorilla was first made known to science through a 
specimen shot by Capt. Oscar von Beringe on Mount Sabinio about 
1902 or 1903 (Matschie, 1903a, p. 253). 

In 1913 and 1914 seven specimens were obtained by E. Arrhenius 
on the volcano Mikeno, Virunga Mountains. "According to Captain 
E. Arrhenius the Gorillas are rather numerous .... They live in 
bands consisting of 20-30 individuals .... 

"The natives hunt the Gorillas to obtain their skin which they 
use for wrapping up their copper thread etc., or for revenging some 
relative. Thus when a man from Sangana had been killed by a 
Gorilla his family killed five Gorillas in revenge. The natives hunt 
Gorillas with the aid of dogs. The dog bites the Gorilla and returns 
to his master who waits for the Gorilla with the spear ready. He 
throws the spear at the Gorilla and runs away. The dog repeats the 
maneuvre, until the animal is killed. The natives do not eat the 
meat of Gorillas, nor that of Chimpanzees." (Lonnberg, 1917, pp. 7, 

Barns (1922, pp. 81-88) encountered a band of Gorillas between 
the volcanoes Mikeno and Karisimbi, and secured a specimen, which 
his hungry native porters refused to eat. "This monster ape would 
seem to have no enemies, failing man; and even man, the most 
dreaded of all the animal world, holds little fear for the gorilla in 
his inaccessible home" (p. 87) . 

"Its food consists, apart from bamboo shoots, entirely of herbage 
docks, sorrels, hemlocks, etc. although honey may be part of the 
menu. He does not grub for roots, neither does he eat fruit as a 
general rule .... 

"Savage man, through superstition as much as anything else, 
but also on account of the inaccessibility of the gorilla's mountain 
home, has left this ape unmolested; we therefore find him and his 
family habitually and fearlessly sleeping on the ground." (Barns, 
1923, pp. 129-130.) 

"As regards longevity, gorillas, on account of their life free from 
molestation, famine, or disease, and also judging by the worn teeth 


of one animal I secured, live, in my opinion, to be a much greater 
age than man" (Barns, 1923, p. 132). 

"The natives of this region have no fear of the gorilla. . . . Some 
of my guides and my gun bearer were trappers and hunters in the 
gorilla forests and were thoroughly familiar with them. At no time 
did the guides or gun boys show any indication of anything more 
than casual interest even when we approached very close to 
gorillas." (Akeley, 1923a, pp. 438-439.) Akeley continues (p. 447) : 

After my first expedition into the gorilla country, I am more convinced 
than ever not only that the gorilla is one of the most fascinating and impor- 
tant objects of study in the realm of natural history, but also that his dis- 
position is such as to permit the most intimate observation of his habits. 
... A few weeks of casual acquaintance and one is fired with a desire to 
ferret out the answers to a hundred questions about this little-known relative 
of man questions of increasing importance to scientists and physicians in 
their efforts to understand and aid man himself. Probably no other project 
of so moderate a size is likely to lead to such immediate and valuable scien- 
tific results as that which will make of the Kivu region a sanctuary, where 
the gorillas under the protection of man may grow more and more accustomed 
to human beings and where through a series of years they may be observed 
and studied. 

On the three mountains, Mikeno, Karisimbi, and Visoke, "I judge 
that there are between fifty and one hundred animals altogether" 
(Akeley, 1923b ; p. 248). 

Akeley's efforts led to the establishment of the Pare National 
Albert, comprising the Kivu volcanoes and providing for the special 
protection of the Mountain Gorillas. Meanwhile Burbridge had 
estimated their numbers at 1,000 to 2,000 individuals. Their range 
extends beyond the volcano region to the bamboo forests dominating 
the highlands of the Great Lakes. (Leplae, 1925, pp. 15, 19.) 

According to Derscheid (1928, pp. 154-159), the animals are 
especially numerous at elevations between 2,700 and 3,500 m., with 
extreme occurrences at 1,900 and 3,900 m. He has met with a few 
solitary old males, but more usually with bands of 7 to 43 indi- 
viduals. He estimates the number on the central massif (Mikeno- 
Karisimbi-Bishoke) at 350 to 500; on the eastern massif (southern 
slopes of Muhabura, Sebyinyo, and Mugahinga) at 150 to 200; and 
in the Uganda portion of the region (northern slopes of the three 
volcanoes just mentioned) at 100 to 150. He also remarks on the 
surprisingly small proportion of young animals among the Gorillas 

"During our two months stay among the different peaks of the 
Birunga Range we observed several herds of Gorillas. The largest 
of these herds consisted of about 20-30 individuals .... In all 
about 70 examples were seen by the members of the Expedition. 
. . . Their stronghold seems ... to be the mountain triangle com- 
posed of Mikeno, Karissimbi and Vissoke. They are mostly found 


in the Bamboo Region, but they also live higher up the steep moun- 
tain slopes with their beautiful vegetation of Hagenia-trees." (Gyl- 
denstolpe, 1928, p. 23.) 

"There is a spotted menace, a potent factor too, in the leopard, 
who destroys numbers of young animals" (Burbridge, 1928, as 
quoted in Yerkes and Yerkes, 1929, p. 398) . 

Pitman (1935, pp. 477-494) gives an excellent account of Gorillas 
in Uganda, and the following excerpts are taken from his paper: 

The occurrence of Gorillas in the Kayonsa region of Uganda [about midway 
between the Birunga volcanoes and Lake Edward] has been known for many 
years (p. 477). 

There is in the Kayonsa a complete absence of bamboo, wild celery, dock, 
and similar juicy-stemmed plants such as abound in the humid, high alti- 
tudes, forcing the Gorilla to confine its diet to a mixture of leaves, berries, 
ferns, the tender fronds of tree-ferns, parts of the wild banana stems, and 
leaves, and fibrous bark peeled off a variety of shrubs in the undergrowth. . . . 

Owing to a lack of what apparently are normal food constituents the 
Gorilla has become more enterprising in search of food, and in consequence 
climbs trees freely to a known height of at least 50 feet. (P. 478.) 

The "beds" of the Kayonsa Gorilla are large platforms built in the trees, 
and often at a considerable height above the ground. 

[The altitude of the habitat varies between 6000 and 7900 feet.] (P. 479.) 

The forest region to the east of the Kishasha river [where some Gorillas 
are known to occur] is a gazetted forest reserve and, in consequence, not 
open for human settlement. There is little likelihood in the immediate 
future of serious conflict between Man and Gorilla in the dense uninhabitable 
valleys to the west of this river and in the vicinity of the Belgian Congo 
border .... 

It was calculated that this western area harboured forty to fifty Gorillas. 
[In the entire region there were possibly at least eighty.] 

Normally the troops vary in size from five to eight or nine, [but one troop 
was said to include nearly two dozen]. (P. 480.) 

The [Kayonsa] Gorilla normally is peaceably disposed and not aggressive 
(p. 483). 

The Wambutte [Pygmies] are extremely tolerant of the Gorillas, but not 
so the other local natives, who would readily endeavour to exterminate the 
lot, were it not for the fact, of which they are well aware, that these splendid 
animals are absolutely protected (p. 484). 

The animals are said to sometimes raid the native gardens but 
not to attack the natives. 

The Chimpanzees 

In spite of the multiplicity of names that have been applied to 
the Chimpanzees, it seems probable that only four valid forms are 
recognizable, representing probably two distinct species, as follows: 

Common Chimpanzee 


Simla troglodytes Blumenbach, Handb. der Naturgesch., p. 65, 1799. ("An- 


Long-haired Chimpanzee; Eastern Chimpanzee 


Troglodytes schweinjurthii Giglioli, Ann. Mus. Civ. Stor. Nat. Geneva, vol. 3, 
p. 114, footnote, 1872. (Upper Uele drainage, Niam-niam country, eastern 
Congo Beige.) 

Western Chimpanzee 


Pan satyrus vents Schwarz, Ann. Mag. Nat. Hist., ser. 10, vol. 13, p. 578, 
June, 1934. ("Sanda Magbolonto chiefdom, Karima district, Sierra Leone.") 

SYNONYMS: For list of synonyms of these three races, see G. M. Allen (1939b, 
pp. 172-175). 

FIGS.: Elliot, 1913, vol. 3, pis. 7, 8, 8 bis (animal); pis. 36-39 (skulls); Yerkes 
and Yerkes, 1929, figs. 69-118. 

Yerkes and Yerkes (1929) write that the "description of the con- 
figuration of the type chimpanzee is as difficult as description of 
man, so numerous and pronounced are individual, sex, and species 
differences and developmental changes." In general, of anthropoid 
form, the forelimbs proportionately long, reaching below the knee 
when the animal stands erect. Form stocky, shape of ear much as in 
Homo, forehead heavily ridged, nose flattened. Face usually bare 
or nearly so, and in the adult black like the skin of the body, except 
in the race verus, in which it is paler. Hair of the head directed 
backward in the typical race, but usually with a parting in verus. 
In the eastern race, schweinjurthii, the hair is longer than in the 
others. The maximum (standing) height of the male is about 5 
feet, of the female 4 feet. Weight of male 125 to 175 pounds; of 
female 100 to 150 pounds. The skull is distinguished readily from 
that of a Gorilla by the smaller teeth and by the fact that when 
viewed from in front the summit of the brain case is visible above 
the brow ridges instead of being hidden by them. Color usually 
black, with often a whitish pygal patch. 

Throughout the vast extent of the tropical rain forest from the 
Gambia and adjacent French Equatorial Africa, south to the 
Congo, and eastward to the borders of Uganda and Tanganyika, 
Chimpanzees are found, but they vary greatly in local abundance. 
They seem much given to wandering about over circumscribed 
areas, and so it is difficult to make censuses or to estimate popula- 
tions. Moreover, the nature of their habitat in rain forest of dense 
growth makes their observation uncertain. Thus in our journey 
across Liberia in 1926, a country in which they are believed to be 
rather common, I saw none, and H. J. Coolidge, Jr., came upon them 
but once in the eastern border of the country. Yet they are com- 
mon in the region about Kindia, in French Guinea, and occur in 


numbers in Sierra Leone and in the forests of the Belgian Congo and 
in the Cameroons. The original specimen was said to have come 
from "Angola" but probably was not native there, for the larger 
species is not now known from south of the Congo. If it was actually 
brought from there, it was no doubt purchased of natives who had 
captured it as a young animal farther north. 

The Chimpanzee offers no trophy for the sportsman and should 
not be killed or captured except for scientific purposes. It is thus 
included in Schedule B of the London Convention of 1933. Its natural 
enemies must be few and, except for man, probably include only the 
Leopard. Native peoples seldom molest them, except where there is 
inducement from whites to capture them for "pets" or to secure 
specimens. Many tribes believe that "every chimpanzee is linked 
with the soul of a man, so that if one is killed the man too will die," 
or some other calamity will ensue. At Kindia in French Guinea, the 
Pasteur Institute maintains a laboratory for the observation and 
medical study of these animals, where individuals may be accus- 
tomed to captivity before being sent to institutions in Europe or 
elsewhere. "From the medical point of view, we have no need of 
emphasizing the advantage to be derived to-day from anthropoids, 
and especially the Chimpanzee, in the study of human diseases ; the 
experimental inoculations of serums, vaccines, and medications of 
all sorts, find in the Chimpanzee a very valuable subject" (Lavau- 
den, 1933, pp. 30-31). Psychological studies of this animal have 
already thrown much light on the evolution of intelligent behavior; 
for a review of such work the reader is referred to the volume by 
Yerkes and Yerkes (1929). 

While the reports of comparative abundance, as noted by travelers 
or persons stationed in parts of its range, are of only relative value 
as often recording mere casual impressions, nevertheless the following 
notes are here added as providing a brief survey of its occurrence 
in selected stations. In the Gambia it is said no longer to exist near 
the coastal towns, but according to E. Johnson (1937, p. 62) every 
year "about fifty animals are brought in for sale from Futa Jalon, 
some 70 miles southeast of Fatoto, 280 miles from Bathurst." They 
are found in the Gola Forest Reserve of Sierra Leone, and small 
troops may be met with by good fortune in the great forests of 
Liberia. On the Gold Coast, according to Haywood (1933), "they 
are only reported from the Western Forest belt, but it seems quite 
possible they are spread over a large area, although by no means 
in large numbers." The Director of Agriculture of the Gold Coast 
writes (in litt., 1937) that the "chimpanzee is now rare and confined 
to the extreme western border of the forest country, but whether 
it was ever plentiful is not known." In Nigeria it is reported from 
forest regions of Oyo, Onitaha, Owerri, Ijebu Ode, and Abeokuta 


Provinces, so probably is present in Benin, Ondo, Calabar, and Warri 
Provinces (Haywood, 1932) . It is apparently common in the Gabun, 
and in the southern Cameroons. It is "numerous" in the equatorial 
forests of the Belgian Congo but in the Ubangi-Shari district is found 
only in the Ubangi Basin, in small numbers (at most a few hun- 
dreds), localized in Haute-Sangha, Lobaye, Ouaka, and Haut- 
Mbomou. It does not seem to have diminished except in the Ouaka. 
In this region it was completely protected since 1916, then partly in 
1931, and once more completely in 1936. It occurs also in the Pare 
National d'Odzala in the Middle Congo and in general appears to be 
threatened not with extermination but with diminution (L. Blancou, 
in letter of 1937) . A. J. Jobaert, in response to queries, writes that in 
the Belgian Congo very few are now killed by Europeans, "but cer- 
tainly the natives, and especially the pygmies, destroy a considerable 
number, although it is totally protected by law ; they were certainly 
quite numerous a few years ago." 

In the eastern part of its range, the race schweinfurthii is locally 
common as far as the border of the rain-forest area in Uganda and 
extreme western Tanganyika. In the upper Congo region, Lang and 
Chapin found it common, as about Aba and Faradje on the north- 
eastern border of the rain forest, and at Avakubi, Niapu, and Medje 
within the forest. On the other hand, reports from the Uganda 
Game Department (1928) indicate that in Kigezi "it would appear 
that the numbers of this species are diminishing. It is, however, 
likely that a permanent change of quarters has resulted in its dis- 
appearance from localities where it was previously known. It has 
been ascertained from Ankole that the parties or families of Chim- 
panzees are great wanderers and not confined to specified localities. 
However, information both from Toro and Bunyoro districts also 
records a recession from areas in which till recently these animals 
were seen and heard." In Tanganyika, Chimpanzees have been 
recorded as far south as the eastern shore of Lake Tanganyika, 
south of Kigoma, in the Mahare Mountains. Footprints and about 
a dozen sleeping platforms were noted by B. W. Savory. They have 
also been found on the west side of the lake as far south as the 
Marungu district. Mr. Savory found collecting of Chimpanzees very 
difficult here, not only on account of the nature of the country but 
also on account of the superstitious fear of the natives, who believe 
these apes are reincarnations of human beings and that a gun fired 
at one of them will surely burst (Dollman, 19356, pp. 15-16) . On 
Mount Kivu Chimpanzees are said to be found but are extremely 
localized. Derscheid records them from Mount Henu and in the 
bamboo forest south of the Karissimbi Volcanoes. 

While such areas of rain forest as are found on some of the more 
eastern isolated mountains, as Kilimanjaro, Kenya, and Elgon, 


might apparently be suitable for Chimpanzees, and in some future 
time might serve as sanctuaries for transplanted stock, they are at 
present uninhabited by these apes. In the distant past, however, 
they may have sheltered them. There is much evidence that in 
eastern Africa the lowlands and slopes of mountains were anciently 
clad with heavy forest but that native races of man have in the 
course of centuries gradually, by burning and cutting, beaten back 
this forest, and that it has given place to scrubby growth or finally 
to scattered thorn scrub. A similar process is slowly going on both 
within the rain forest and at its edges, with slow but gradual clearing 
of trees for agriculture, and subsequent abandonment. Continued 
long enough, this results in final destruction of the high forest, first 
in spots, then in local areas, and finally over larger tracts, all of 
which will eventually much curtail the available living areas for 

G. M. A. 

Lesser Chimpanzee; Pygmy Chimpanzee 


Pan satyrus paniscus Schwarz, Revue Zool. Africaine, vol. 16, p. 425, April 1, 
1929. (South of the upper Maringa River, 30 km. south of Befale, south 
bank of the Congo, Congo Beige.) 

FIGS.: Coolidge, 1933, pi. 1, figs. A, B; pi. 2, fig. A. 

Present evidence seems to indicate that this is a smaller species 
than the Common Chimpanzee and distinct from it. Its known 
range is in the Middle Congo forests, on the south side of the river, 
here supposed to form a physical barrier to northward extension. 

Coolidge (1933) has summed up our knowledge of this animal and 
has made a comparative study of its skeleton. An adult female 
(containing a fetus) had a head and body length of 630 mm.; 
height from crown to sole, 1,010; spread of arms, 1,510. It is thus 
much smaller than the other species. The hair is fine in texture and 
glossy black throughout except for a small white pygal tuft, and is 
long and dense, without a parting on the head. The skull has a 
rather juvenile appearance in its inflated forehead and small brow 

Although the existence of a Chimpanzee on the south side of the 
Congo had been several times reported, it was not until 1928 that a 
M. Ghesquiere obtained specimens for the Congo Museum in Ter- 
vueren and the animal was described. Previously a specimen had 
been in the British Museum, collected in 1895. Other specimens 
have since been secured, and more information is likely soon to be 
placed on record. Dr. James P. Chapin, who secured a specimen in 
1930 near Lukolela, describes the voice as neither so loud nor so shrill 


as that of the larger Chimpanzee. Dr. R. M. Yerkes had a specimen 
in captivity for over a year in 1923-24 and has written of its 
behavior. Very little is known of its abundance, but one may sup- 
pose its distribution includes the rain-forest area between the 
Congo and the Kasai. 

G. M. A. 

Order EDENTATA: Edentates 

Family MANIDAE: Pangolins 

Three genera of this family, represented by four species, occur 
in Africa south of the Sahara, and all of them are treated in the 
following pages. Two other genera (Manis and Phatages), repre- 
sented by five forms, occur in the Oriental region (India, China, 
Siam, Indo-China, Malaysia) ; while subjected to some perse- 
cution by reason of their supposed medicinal value, these Oriental 
pangolins are not included in the present report. 

Giant Pangolin or Scaly Anteater. Pangolin geant (Fr.). 
Riesenschuppentier (Ger.) 


Manis gigantea Illiger, Abhandl. K. Akad. Wissen. Berlin, physik. Kl., 1804- 

1811, p. 84, 1815. (Guinea = West Africa.) 
FIGS.: Buttikofer, 1890, vol. 2, p. 394, fig.; Beddard, 1902, p. 190, fig. 109; 

Schubotz, 1912, p. 357, fig.; Bequaert, 1922, pi. 24, fig. 2, pi. 25, fig. 2; 

Schouteden, 1930, p. [14], fig. 3a; Halt, 19346, pis. 32-34, and 1934c, 

p. 727, upper fig., p. 729, fig.; Rosevear, 1937, p. 12, fig. 2. 

The various species of African pangolins do not appear to be 
numerous anywhere. They are in considerable demand among the 
natives, and active hunting keeps down their numbers. All forms 
are placed in Schedule B under the London Convention of 1933. 

In all African pangolins "no hairs project between the scales, the 
median dorsal row of scales does not extend to the tail tip, and 
there is no external pinna of the ear." In S. gigantea and S. tem- 
minckii "the belly is naked, the preaxial surface of the fore limb 
bears scales to the base of the claws, the tail is massive and bears 
no naked subterminal pad." In the former "there are 12 to 15 
scales in the median dorsal row of the tail. . . . The scales of the 
head, neck, shoulders, arm, and hind legs are dominantly dark olive- 
brown. This color shades gradually into avellaneous over the dorsal 
region. . . . Over the tail a deep Roman green assumes increasing 
prominence in the apical part of the scale. . . . The species is hair- 
less, except for a dense ring of short, circumorbital bristles and a 
patch of similar hairs in front of the auditory meatus." The tail 


length averages a little less than half of the total length. (Hatt, 
19346, pp. 646-649.) The species attains a total length of 1,710 mm. 
and a tail length of 830 mm. (Allen and Coolidge, 1930, p. 606) . 

The range appears to extend from Sierra Leone and Liberia east 
to the Ubangi-Shari Territory of French Equatorial Africa and 
northeastern Belgian Congo. It corresponds rather closely to the 
Upper and Lower Guinea Forest District of Chapin (1932, p. 90) 
and of Bowen (1933, pp. 256, 258). "M. gigantea is known from the 
West African Rain Forest and the adjoining wooded galleries" (Lang, 
in Bequaert, 1922, p. 325). Matschie (1894a, p. 5) seems to extend 
the range as far as Senegambia. 

FIG. 20. Giant Pangolin or Scaly Anteater (Smutsia gigantea). After 
photo by Lang. 

Sierra Leone. A species of pangolin, said to be Smutsia gigantea, 
is of fairly general distribution but is not commonly seen. It pro- 
vides food for the natives. There is no evidence of depletion, and 
no protective measures are taken. (Colonial Secretary's Office, in 
litt., July, 1937.) 

Liberia. This is a very rare animal in Liberia. A specimen 
secured by Jackson at Cape Mount had consumed a large quantity 
of termites and driver ants. The flesh is very tough and has a flavor 
of formic acid. (Buttikofer, 1890, vol. 2, pp. 395-396.) 

Another specimen is recorded by Jentink (1888, p. 56) from Little 

A male of record size was obtained from natives at Paiata (Allen 
and Coolidge, 1930, p. 606) . 

Gold Coast. This species "is found in the savannah areas of 
N. Ashanti and the Northern Territories. 

"There is little doubt that all [the species of pangolins] are now 
much less common than formerly, though it is probable that their 
range has not decreased. 


"Night hunting and the use of wire snares are the main causes of 
depletion. Their meat is considered one of the greatest delicacies." 
(Assistant Conservator of Forests, Gold Coast, in litt., July 22, 

Nigeria. The species is reported from Nigeria, but without a 
definite locality record (Rosevear, 1937, p. 13) . 

French Cameroons. It occurs in the forest region and is absolutely 
protected except under scientific permit (Paris Agency, in litt., 
November, 1936). 

Gabun. The Giant Pangolin is confined to the great forest. Only 
a skin has come under personal observation. It was, however, 
abundant at Mimongo in the region of Akelai. (A. R. Maclatchy, 
in litt., February 5, 1937.) 

Ubangi-Shari district, French Equatorial Africa. It appears to 
be localized in the forested region. It is not threatened, and has 
been totally protected since 1929. (L. Blancou, in litt., December, 

Belgian Congo. Schubotz (1912, p. 356) records a specimen from 
Angu, on the Uele River. 

Lang (in Bequaert, 1922, p. 320) says of the several local species 
of pangolins: 

"The signs of their fossorial practice are as often a cause of their 
discovery as is the strong odor they emit, and dogs of native hunters 
never fail to challenge their presence. Various highly valued talis- 
mans, which their captors obtain from the claws, scales, hairs, and 
other parts of some of the scaly ant-eaters, suffice to make them an 
always welcome prize and their meat is an additional incentive for 
their destruction." 

Lang also describes (p. 325) a Pygmy method of capturing the 
present species: "Pygmy boys, with one end of a strand of rattan 
fastened to the waist and the other held by friends waiting outside, 
entered the burrows without hesitation .... These boys, armed 
only with a knife, merely fastened the rattan around the live pan- 
golin, which they prodded from behind while their companions 
pulled it out of the hole. These otherwise harmless beasts, when 
touched while rolled up, suddenly switch their tail sidewise with such 
force that, if one's hand is caught between the rough body scales 
and the tail, it is seriously mutilated by the shearing action." 

Lang records (p. 325) specimens from Bafuka, Niangara, Poko, 
and Niapu in northeastern Belgian Congo. 

"The only specimen I was ever able to obtain was dug out for 
me by natives, with the expenditure of much labor and time, on the 
Semliki side of the forest" (Christy, 1924, p. 228). 

Schouteden states (1930, p. [95]) that the species ranges from 


the Lower to the Upper Congo. He also records (1935, p. [62]) a 
specimen from the Kivu region. 

The several species of pangolins do not appear numerous any- 
where in the Belgian Congo. The natives do not hunt them espe- 
cially, and the Europeans not at all. Brush fires alone destroy a 
great many. (A. J. Jobaert, in litt., November 10, 1936.) 

South African Pangolin ; Scaly Anteater. I jzer Magauw ; 
letermago (Boer) 


Mani's temminckii Smuts, Enumeratio Mammalium Capensium, p. 54, pi. 3, 

figs. 6-7, 1832. ("E regionibus, ultra Latakou sitis" = probably the region 

north of Litakun, British Bechuanaland.) 
FIGS.: A. Smith, 1849, pi. 7; Royal Nat. Hist., vol. 3, p. 229, fig., 1894-95; 

Matschie, 1895, p. 143, fig.; W. L. Sclater, 1901, vol. 2, p. 217, fig. 148; 

Fitzsimons, 1920, vol. 4, pi. facing p. 233. 

Though widely distributed in South and East Africa, this seems 
to be a decidedly scarce animal. 

"General form somewhat elongated and lizard-like, covered every- 
where, except on the lower surface of the head and body and inside 
the limbs, with a series of over-lapping broad scales of a dark horn- 
brown colour with paler edges and tips; head very small and 
pointed," covered above with small scales; "no external ear . . . . 
Across the middle of the back eleven rows of scales . . . ; limbs 
short each with five toes and claws .... Tail very broad," cov- 
ered above and below with 4-5 rows of scales. Head and body, 24 
inches; tail, 18. (W. L. Sclater, 1901, vol. 2, pp. 217-218.) 

"The scaly ant-eater is chiefly found to the north of the Orange 
River, though said to occur rarely in Prieska and the other districts 
just south of the river; from here it extends through the Orange 
Free State, the Transvaal, Bechuanaland, the Kalahari and German 
South-west Africa to Rhodesia; north of the Zambesi it occurs in 
South Angola, Nyasaland and East Africa as far as Somaliland." 
(W. L. Sclater, 1901, vol. 2, p. 218.) Matschie (1894, p. 5) extends 
the range north to southern Kordofan (about lat. 17 N.). 

"Well known in the Orange Free State (Ventersburg Albany 
Museum) , the Transvaal, Bechuanaland, Ngamiland, and Southern 
Rhodesia." Also "recorded from Northern Rhodesia, Nyasaland, 
and according to Sclater Uganda, East Africa and Somaliland." 
(Shortridge, 1934, vol. 2, p. 665.) 

Cape Province and British Bechuanaland. At Litakun, British 
Bechuanaland, Burchell (1824, vol. 2, pp. 423-424) observed a skin 
lying on the hedge of a native cattle enclosure, "placed there . . . 
to preserve the cattle from the evil effects of sorcery. . . . When- 


ever a recent track is met with, the animal is traced to its hole and 
dug out if possible, as the flesh, which is extremely fat, is esteemed 
so great a delicacy that the law requires that every khaaka which 
is killed shall be brought to the Chief." 

Referring to this same general region, A. Smith says (1849, text to 
pi. 7) : "Only one solitary specimen of this species was obtained by 
the expedition before reaching 26 south latitude, and but two more 
between that parallel and the tropic of Capricorn .... Its extreme 
scarcity probably arises from its having long been zealously sought 
after by the natives .... Whenever a specimen ... is secured, 
it is immediately burned in some cattle pen, which, according to 
the opinion of the sacrificer, tends to increase the health and fertility 
of all cattle who may afterwards enter the fold. Not many years 
ago a specimen was captured in the northern part of the Cape 

"I have seen a dry skin from Upington " (Shortridge, 1934, vol. 2, 
p. 665) . In Griqualand West the animal is now very scarce, and the 
scales are used by the natives as medicine (McGregor Museum, 
Kimberley, in litt., June, 1937) . 

South-West Africa. In the Omaheke and the Kaukauveld it is 
widespread but rather rare; skins are seen occasionally among the 
natives. It is also reported by Bushmen in the Hukweveld. (Zu- 
kowsky, 1924, p. 68.) 

"The Pangolin occurs throughout South-West Africa," but is 
"never plentiful. . . . 

"It is apparently most numerous north of the Tropic of Capricorn 
and in the sand-plains adjoining Bechuanaland. 

"Rare in the vicinity of the Orange River and in the southern 
parts of Great Namaqualand. . . . 

"Pangolin scales (used as charms) were not infrequently seen in 
the possession of Bushmen and other natives." (Shortridge, 1934, 
vol. 2, p. 665.) 

Angola. Monard (1935, p. 183) records two specimens from the 
region between the upper Kului and the Kubango, where the natives 
report the animal as rather common. Monard also mentions (p. 185) 
specimens recorded by Bocage from Caconda and Mossamedes. 

Transvaal. "A number of examples have been sent to the National 
Zoological Gardens, chiefly from the Rustenburg and Marico districts 
of the Transvaal" (Haagner, 1920, p. 237). 

"ISIever very common and probably scarcer now as a result of 
closer settlement. There is a demand for its scales by native witch 
doctors for 'medicine/ as much as six pence per scale being paid, 
so that this leads to a considerable amount of destruction of the 
animal. Being entirely useful and harmless it should receive more 
protection than is actually accorded it .... (Not included in the 


game laws, i. e., without any special protection.) " (A. Roberts, in 
Hit., November, 1936.) 

Portuguese East Africa. Peters (1852, p. 174) records specimens 
from Quitangonha, from near Cape Delgado, and from the vicinity 
of Quelimane. He adds that the scales are made into finger rings 
and worn as a protection against the "evil eye." 

Kirk states (1865, p. 654) that it occurs near Sena. 

Southern Rhodesia. Chubb (1909, p. 125) records a specimen 
from AVankie, Matabelelend. 

The species is by no means common. Most Rhodesian natives use 
the skin as a charm, and for this reason the animals are in great 
demand. Were it not for their retiring nature and strictly nocturnal 
habits, they would be in danger of extermination. They will be 
protected in the near future. (Game Warden, Wankie Game Re- 
serve, in litt., March, 1937.) 

Northern Rhodesia. Pitman notes (1934, p. 173) that this pango- 
lin is "recorded from Batoka Province and Barotse." The natives 
do not "seem to know of it in the areas I have toured (with the 
exception of the Kafue Hook) ." He quotes Neave (1906) to the effect 
that it is not unusual to see the scales worn as charms by natives 
of the mid-Zambesi Valley. 

Tanganyika Territory. Holmwood (1878, p. 632) records a speci- 
men "from the coast opposite Zanzibar, lat. 6 S. ; but I have seen 
what I took to be the same animal, both in Somali-land under the 
equator and as far south as the Makna country opposite Mozam- 

Matschie (1895, p. 143) records the animal from Wahumba, 
Bagamoyo, Massai Nyika, and Mandera. 

The Game Preservation Department (in litt., December, 1936) 
reports no danger of extinction. 

Kenya. The Game Warden (in litt., November, 1936) reports 
no decrease, though the animal is not protected. 

Three-cusped Pangolin; White-bellied Pangolin; Pale-bellied 

Pangolin; Pointed-scaled Pangolin. Pangolin tricuspide 

(Fr.). Dreizackige Schuppentier (Ger.) 


Manis tricuspis Rafinesque, Annales Gen. Sci. Physiques [Bruxelles], vol. 7, 
p. 215, 1821. (Type locality not stated; restricted by Allen and Coolidge 
(1930, vol. 2, p. 606) to "West Africa.") 

FIGS.: Royal Nat. Hist., vol. 3, p. 230, fig., 1894-95; Johnston, 1906, vol. 2, 
p. 749, fig. 292; Schouteden, 1930, p. [88], fig. 1; Hatt, 19346, pis. 36-37, 
and 1934c, p. 727, lower fig., p. 730, upper fig., p. 731, right-hand fig.; 
Rosevear, 1937, p. 12, fig. 1. 


This species occurs in apparently larger numbers than the other 
African pangolins. 

It is "an arboreal species with a tail constituting over half the 
total length. The characters of the tail tip and the fore limbs are 
like those of Manis longicaudatus. The scales, however, are small 
and numerous, brown, and during mid-life, tricuspid. The post- 
scapulars are not enlarged." Under parts grayish white. In young 
animals "the margins of the scales are even, but with ensuing wear 
. . . the scales become sharply dentate, or, usually later, tridentate." 
In old age the animals have "cuspless, worn, elongate scales." In 
half-grown and mature animals the unsealed parts of the skin are 
covered with hair, attaining a length of 20 mm. Longitudinal rows 
of scales, 21-25; marginal caudal scales, 35-40. Total length, up to 
1,027 mm.; tail, 607 mm. (Hatt, 19346, pp. 655-658.) 

This pangolin is not confined to the Upper and Lower Guinea 
Forest Districts but ranges southward into the Southern Congo 
Savanna District and eastward into the Uganda-Unyoro Savanna 
District of Chapin (1932, p. 90) and of Bowen (1933, pp. 256, 258). 
Hatt (1934b, p. 656) records specimens from Liberia, the Ivory 
Coast, Cameroons, Fernando Po, Gabun, the lower Congo, Kasai 
district, and central Angola. According to Matschie (1894a, p. 6), 
the range extends west to Gambia, and Jentink (1882, p. 208) has 
a record from Sierra Leone. 

Liberia. The species appears to be distributed over the entire 
region. Specimens are recorded from Buluma, Schieffelinsville, 
Junk River, Hill-town, and Farmington River. The animal can be 
tamed and kept a long time in houses, where it runs free and preys 
upon ants, cockroaches, and other troublesome insects. (Buttikofer, 
in Jentink, 1888, p. 57.) 

Allen and Coolidge (1930, vol. 2, p. 606) record "a native-made 
skin bought at Since." 

Gold Coast. Hayman (1936, p. 937) records specimens from 
Goaso and Mampong. 

The species is found through much of the forest country, but is 
doubtless much less common now than formerly (Assistant Con- 
servator of Forests, Gold Coast, in litt., July 22, 1937) . 

Fernando Po. Fraser (1848, text to pi. 28) records the species 
from this island, where "the flesh is said to be exceeding good eating, 
and is in great request among the natives." 

Gabun. This pangolin is confined to the great forest. Although 
legally protected, it is actively hunted by the forest natives, who 
capture great quantities. To prevent this is difficult, for the animal 
is taken in trigger traps set for small game. The real safeguard 
would be the prohibition of this type of trap; but those who know 


the brush know how much such prohibition would be worth. (A. R. 
Maclatchy, in litt., February 5, 1937.) 

French Equatorial Africa. Matschie (1894a, p. 6) records the 
species from Loango. 

It is common almost everywhere in the Ubangi-Shari district. 
It has been totally protected since 1929, and is not threatened. (L. 
Blancou, in litt., December, 1936.) 

Angola. Monard (1935, p. 185) quotes Bocage to the effect that 
this pangolin is rather common at Bembe and Malange; he also 
gives records for Bimbi and Cazengo. 

Belgian Congo. Schwarz (19206, p. 1061) records specimens 
from Libenge on the Ubangi, Panga on the Aruwimi, Angu on 
the Uele, and Avakubi on the Ituri; also from Kudurma and Ka- 
bayendi in the Niam-Niam country (not far from the Congo-Sudan 
boundary) . 

Lang (in Bequaert, 1922, pp. 320-323) remarks that tricuspis 
is the commonest of the African pangolins. "Being timid, they 
readily make use of their natural safeguard and, when even slightly 
annoyed, roll up in a ball .... When forcibly unrolled, they 
may succeed in driving off their tormentors by well directed jets of 
an ill-smelling, acrid liquid from the anal region; native dogs suffer 
for a considerable time from the effect of this substance, which 
greatly irritates their mucous membranes. . . . 

"If unmolested and placed near their favored prey, they uncoil 
readily .... One soon realizes how thoroughly they are special- 
ized as ant-eaters, for their methods of attack and disposal of ants 
are as effective as their ways of guarding themselves against the 
defensive means of their prey. In the regions we visited, the pan- 
golins preferred true ants, as stomach contents clearly showed, 
though many of our captives would plunder termitaria with great 
eagerness. . . . 

"One taken near a column of army ants (Dorylus) merely made 
good its escape, another quickly broke up the well-ordered line. 
. . . Lashing its sticky tongue through the confused crowds, the 
ant-eater lost no time in moving back and forth along the ant 
column as quickly as the dense clusters vanished into its mouth. 
Its hunger satisfied, it at once retreated, freeing itself of the few 
army ants that had managed to dig their mandibles into the soft 
parts of its hide. M. tricuspis fed freely on many other kinds of 
ants. Those we had alive at Avakubi, Medje, and Niapu were 
particularly fond of ants of the genus Myrmicaria. . . . 

"African pangolins have helped to enrich the stores of witchcraft." 

Hatt (19346, p. 645) records 66 specimens from Akenge, Avakubi, 
Faradje, Gamangui, Medje, Ngayu, Niangara, Niapu, Poko, and 


" Uganda. An arboreal pangolin (presumably tricuspis) is reported 
by Johnston (1902, vol. 1, pp. 395-396). 

"Two or three species occur in Uganda, the common representa- 
tive being Phataginus tricuspis, a forest species. There is no reason 
to believe that Pangolins are any less plentiful than formerly. In the 
Mabira Forest P. tricuspis is abundant. All species of Pangolins 
are completely protected in Uganda." (Game Warden, Uganda, 
in litt., December, 1936.) 

Long-tailed Pangolin; Black-bellied Pangolin. Pangolin a 

longue queue (Fr.). Langschwanzige 

Schuppentier (Ger.) 


Pholidotus longicaudatus Brisson, Regne animal, vol. 3, Quadr., p. 19, 1762. 

("Probably West Africa" (Allen and Coolidge, 1930, p. 606).) 
SYNONYMS: Manis teiradactyla Linnaeus (1766); Manis macroura Erxleben 

(1777); Manis hessi Noack (1889). 
FIGS.: Noack, 1889a, pi. 1; Johnston, 1906, vol. 2, p. 753, fig. 295; Bequaert, 

1922, pi. 25, fig. 1; Allen and Coolidge, 1930, pp. 603-605, figs. 447-449; 

Schouteden, 1930, p. [94], fig. 3a; Halt, 19346, pi. 35, figs. 1-2, and 1934c, 

pp. 726 (both figs.) and 731 (lower fig.) ; Rosevear, 1937, p. 12, fig. 3. 

The very limited amount of information available concerning 
this species suggests that it is one of the rarest of the African 

This is "an arboreal species with a long prehensile tail, equaling 
about two-thirds of the total length. . . . The forearms bear no 
scales, but are covered with hair. The scales are large, yellow, and 
on the flanks are keeled. The two inferior postscapular scales are 
markedly larger than those adjacent to them. . . . The belly hair 
is black in most individuals .... The whole face . . . dark brown, 
nearly black." Total length up to 937 mm.; tail, 645. (Hatt, 19346, 
pp. 651-652.) Thirteen rows of scales on the body; 44 marginal 
scales on the tail; two rows of 9-10 scales before the tail tip (Mat- 
schie, 1894a, p. 7). 

The range appears to be more or less coextensive with the Upper 
and Lower Guinea Forest Districts of Chapin (1932, p. 90) and of 
Bowen (1933, pp. 256, 258). Jentink (1882, p. 207) records speci- 
mens from as far west as Senegal and Sierra Leone. Otherwise the 
species is known from Liberia to Gabun and the northeastern Bel- 
gian Congo. 

Liberia. The animal is pretty rare, though a number of living 
specimens were received, including one at Soforeh Place. (Butti- 
kofer, 1890, vol. 2, pp. 393-394.) 

Jentink (1888, p. 56) records additional specimens from Hill-town 
and Farmington River. 


Live specimens were brought to Allen and Coolidge (1930, vol. 2, 
p. 606) at Lenga Town on the Farmington River and at Paiata. 

Gold Coast. Specimens are recorded from Dabocrom and Elmina 
(Jentink, 1882, p. 207) ; also from Goaso (Hayman, 1936, p. 937). 

The species is found through much of the forest country, but there 
is little doubt that it is now much less common than formerly 
(Assistant Conservator of Forests, Gold Coast, in litt., July 22, 

Cameroons. Hatt (19346, p. 653) records the species from this 
country, without stating the exact locality. 

Gabun. Hatt (19346, p. 652) records a specimen from Fernand 

Belgian Congo. Noack (1889a, p. 100) based his name Manis 
hessi upon a specimen from the vicinity of Banana, at the mouth 
of the Congo. 

Hatt (1934b, pp. 651, 653) records specimens from Bolobo and 
Lukolela on the Lower Congo, and from Akenge, Gamangui, Medje, 
and Niapu in the northeastern part of the country. 

Order RODENTIA: Rodents 

Family LEPORIDAE: Hares and Rabbits 

This family is of nearly cosmopolitan distribution; but it is 
absent from Madagascar and part of the Malay Archipelago, and it 
was lacking in Australia until introduced. There are about 11 genera 
and over 200 species and subspecies. There is generally an abun- 
dance of individuals, and only a single species, the insular Amami 
Hare, comes within the scope of this report. 

Amami Hare; Liu Kiu Hare 


Caprolagus furnessi Stone, Proc. Acad. Nat. Sci. Philadelphia 1900, p. 460, 
1900. ("Liu Kiu Islands.") 

This remarkable insular hare, unique representative of its genus, 
has been suitably recognized and protected by the Japanese Govern- 
ment as a "Natural Monument." 

Size approximately that of Lepus americanus; hind foot, tail, 
and ears remarkably short ; claws very large and strong ; soft under- 
fur plumbeous ; long hairs coarse and hispid, brownish black, many 
with buff annulations; a median black stripe from neck to rump; 
under parts mostly pale buff. Total length of flat skin, 550 mm.; 
tail, about 8 mm., (Stone, 1900, pp. 460-461.) "Pentalagus is the 
most marked of any of the genera of the Leporidae, the tooth 


formula, the structure of the teeth, the relative size of the radius and 
ulna, and the very short tarsus and metatarsus being peculiar to 
the genus and unlike anything in the rest of the family" (Lyon, 1904, 
p. 430). 

In the original description Stone (1900, p. 460) records two speci- 

Thomas (1906a, p. 357) records a specimen from "Oshima, Oki- 
nawa, Liu-Kiu Is.," and adds: "Another specimen is now living in 
the Duke of Bedford's menagerie at Woburn." 

"The distribution of this species ... is restricted to the Islands 
of Amami-Oshima and Tokuno-shima in the Loochoo archipelago 
where it is endemic" (Kaburaki, 1934, p. 4183) . 

"Number is unknown, but as it is carefully protected as one of the 
'Natural Monuments/ by the Law for Preserving Scenery, Historic 
and Natural Monuments, and it is also strictly prohibited to capture 
the species without special permission, and besides it is forbidden 
by the game law, it will never become extinct" (Nagamichi Kuroda, 
in litt., July 5, 1938) . 

Family CASTORIDAE: Beavers 

The single genus of this family is repres'ented by one species 
(Castor canadensis) , with 20 subspecies, in North America, and by 
another species (fiber) , with perhaps four subspecies, in Europe and 
northern Asia. All the American forms have been treated by Dr. 
Allen in the preceding volume (1942), and an account of Castor 
fiber and its subspecies follows here. It is primarily the demands 
of the fur trade that have brought about trie deterioration in the 
status of the Beavers. 

European Beaver. Castor; Bievre (Fr.) Biber (Ger.) 


[Castor] fiber Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 58, 1758. (Sweden.) 
FIGS.: Geoffrey and Cuvier, Hist. Nat. Mamm., vol. 6, pi. '275, 1824; Brandt 
and Ratzeburg, 1829, pi. 3; Blasius, 1857, p. 403, fig. 224; Royal Nat. Hist., 
vol. 3, p. 97, pi., 1894-95; Collett, 1898, pi. 12; Martin, 1910, pi. 10; 
Didier and Rode, 1935, p. 188, fig. 98. 

While treated here as a specific unit, Castor fiber has been divided 
into a number of forms, including the following which are con- 
sidered by Kuntze (1935, p. 64) to be more or less tenable: 

C. /. fiber Linnaeus (Sweden) ; 

C. f. vistulanus Matschie (western Poland) ; 

C. /. albicus Matschie (Dessau, Anhalt, Germany) ; and 

C. /. galliae Geoffrey (the Rhone, France). 


The former range of the Old World Beaver included the forested 
regions of Europe and northern Asia. The original colonies are now 
extinct in all save a few localities in France, Germany, Norway, 
Poland, Russia, and Siberia. In recent times the animal has been 
reintroduced into England, Sweden, and Latvia. 

The general form is heavy and thickset; eyes and ears small; 
hind feet large, broad, and webbed, the claw of the fourth digit 
with a horny, compressed supplement; tail scaly, mostly naked, 
broad, depressed ; general color a peculiar and very uniform clayey 
buff, the under parts a little more yellowish. Head and body (fe- 
male) , 820 mm. ; tail, 380 mm. (Miller, 1912, pp. 948-952.) Weight, 
15 to 25 or even 36 kilograms (Didier and Rode, 1935, p. 188). 

Great Britain. In Wales, in A. D. 940, Beaver hides were req- 
uisitioned for making the borders of the king's garments; it was 
evidently then a rare animal. In 1188 it was still found on at least 
one river in Wales and on a single river in Scotland, though it had 
apparently died out quite generally in other parts of Great Britain. 
"The written records we have of its occurrence are very frag- 
mentary, and not wholly satisfactory." Remains have been exhumed 
in both England and Scotland. Various place names in England 
indicate the former occurrence of Beavers there. (Harting, 1880, 
pp. 33-46.) 

Their skins were exported from England and Scotland until the 
middle of the twelfth century. Beavers x were reintroduced on the 
island of Bute, Scotland, in 1874, but died out about 1890. There 
were similar introductions in Suffolk, England, in 1870, and in Sussex 
at some time prior to 1905. (Millais, 1905, pp. 162-163.) 

In 1663 a good Beaver hat in England cost 85 RM. in German 
currency (Kriiger, 1931, p. 54). 

Spain. Strabo, writing of this country in the first century B. C., 
is said to mention the Beaver as a well-known animal (Blasius, 1857, 
p. 407; Kriiger, 1931, p. 52). 

France. The Beaver was once widely distributed in France, 
being found on many watercourses in various basins. It gradually 
became rare, but in the sixteenth century was still found on many 
rivers, principally the Oise, the Somme, and the Marne. Today it 
is found only on the Rhone and its tributaries, below Valence. The 
principal habitats are: (1) the mouth of the Ardeche; (2) the 
mouths of certain small watercourses the Ceze, the Tave, and the 
Aigues; (3) the vicinity of Roquemaure and the lie de Miemas; 
(4) the vicinity of Avignon and the lie de Barthelane; (5) along the 
course of the Garden; (6) between Tarascon and Beaucaire; (7) on 

i According to Kriiger (1931, p. 53), Scottish importations at this period were 
of Canadian Beavers. 


most of the course of the Petit-Rhone, in rather numerous colonies ; 
(8) on the Grand Rhone, beginning at Aries, in less numerous 

The 60 or 70 known stations are certainly not the only ones. A 
rough estimate of the total population is 300 individuals. 

The reasons for depletion are numerous and diverse. Although 
the Beaver was always hunted for its valuable fur, it was long 
considered, up to recent years, as harmful and thus was under official 
ban. Trapping in submerged nets has been particularly fatal. 
The frequent floods on the Rhone have been a serious factor in 

FIG. 21. European Beaver (Castor fiber). After Brehm. 

Perhaps pollution of the watercourses in certain areas is to be 
blamed, for autopsies have revealed tubercular lesions. It does not 
appear from the autopsies, however, that the fecundity of the species 
has been diminished. 

When its existence was threatened a dozen years ago, the warning 
issued by certain naturalists rapidly bore fruit. Restrictions on hunt- 
ing, establishment of reserves, warden service, propaganda in favor 
of the Beaver, and appreciation of this rare animal by the local 
population, have been effective in its conservation. Its. future seems 
brighter, and in general the colonies seem more prosperous than a 
little while ago. It would be easy to improve the situation by the 
creation of more reserves, by the establishment of zones of refuge 
at the time of floods, and by the repopulation, if possible, of old 
abandoned colonies. (E. Bourdelle, in litt., March 6, 1937.) 

Since 1909 the hunting and capture of Beavers have been pro- 
hibited for all time. Twenty kilometers of the Rhone have been 


declared a protected reserve. As a result of persecution the species 
has lost the habit of constructing dams and lives in burrows in the 
banks of the streams. (Didier and Rode, 1935, pp. 192-193.) 

Trouessart wrote in 1884 (pp. 119-121) that its northern limit 
in the Rhone Basin was approximately Valence (as it is now). He 
added that it was becoming rarer each year, owing to relentless per- 
secution. Also that its flesh is excellent, and since it is hunted for 
its hide and for its castor, as well as because of its depredations in 
young plantations, its early exterminaton in France could be pre- 
dicted. But such a fate has been happily warded off. 

A few years ago its castor was worth more than 250 francs a 
pound. For a long time the Syndicat des digues du Rhone paid a 
bounty of 15 francs on Beavers, on account of the alleged damage 
to dikes. But with better information the bounty was abandoned. 
(Martin, 1910, p. lOb.) 

Italy. Gesner (1551, vol. 1, p. 337) mentions the Beaver as 
occurring at the mouth of the Po. 

Yugoslavia. The species is entirely exterminated, the last speci- 
mens having been observed in 1859 at Syrmia on the Danube and 
at about the same time in Bosnia on the River Ukrina. A good 
many fossil remains have been found in Croatia and Slovenia. (M. 
Hirtz, in Hit., December, 1936.) 

Rumania. The species existed in Transylvania up to about 1500, 
and in Moldavia to 1823 (Calinescu, 1931, p. 82). 

Hungary. Extermination took place in the first half of the nine- 
teenth century (J. Schenk, in litt., November, 1936). 

Austria. In 1867 Beavers still lived northwest of Salzburg, where 
the Sur discharges into the Salzach, but by 1870 only disused 
burrows could be found. There was formerly a protected colony 
in the plains of the Danube near Wien. But now the Beaver is no 
more to be found in the Danube region. In 1861 the castor fetched 
600 Gulden in Salzburg. (Kruger, 1931, pp. 52-54.) 

Czechoslovakia. Under the protection of the Princes of Schwarz- 
enberg, Beavers survived long in the tributaries of the Moldau, but 
the last of them died in 1883 (Kruger, 1931, p. 53). 

Switzerland. Gesner (1551, vol. 1, p. 337) reports the Beaver 
as a very common animal in the large rivers. But it could not sur- 
vive strong persecution (Kruger, 1931, p. 52). Millais (1905, p. 160) 
quotes Harting to the effect that "Beavers were to be found in the 
Aar, the Linnet, and the Reuss, and up to the last century [eight- 
eenth] a few still lingered on the banks of the last-named stream, 
on the Thiele, and the Byrse." 

Germany. On the Rhine the animals died out more than 300 
years ago. In Westphalia they occurred up to the middle of the 
nineteenth century, and the very last was killed apparently in 1877. 


In the Province of Hannover they disappeared more than a hundred 
years ago. 

Beavers now find refuge only on the middle course of the Elbe 
between Wittenberg and Magdeburg, together with its tributaries 
the Mulde, Saale, and Nuthe, and the adjacent Altwasser. The 
population was estimated in 1890 at 200; in 1913, at 188; in 1919, at 
42; in 1922, at 200; in 1926, at 164; in 1929, at 263. 

The almost total destruction of the Beaver in Europe is to be 
explained only by avaricious persecution. It was pursued because of 
its tasty flesh, its valuable pelt, and especially its castor, which 
commanded a very high price as a panacea. 

The presence of the Beaver today on the middle Elbe is due to 
certain protective measures. Formerly it enjoyed no protection. 
The Prussian Game Law of 1907 gave it a 10-month closed season. 
In 1921 and again in 1929 it was given complete protection. Along 
the Elbe mounds are constructed to furnish a refuge during floods, 
and some willow plantations are provided as food. There are re- 
strictions on fishing and trapping in the immediate vicinity of the 
Beaver burrows. The Provincial Assembly has made an appro- 
priation of 1,000 RM. for settlement of claims for damage by 
Beavers. Despite these protective measures, the Beaver stand in- 
creases only slightly or not at all. 

Owing to the penalties involved and the difficulty of disposing 
of the skin, deliberate killing for profit has practically ceased. 
But some animals still fall victims each year to the human lust for 
killing. More serious is the killing for protection against damage. 
In the eyes of the country people and the fishermen the Beaver is 
injurious. It steals their potatoes and turnips and destroys their 
fruit and forest trees. Some are caught and drowned in fish nets 
and traps. They are also endangered by steel traps set for Otters. 

Floods and drifting ice constitute the greatest menace to the 
Beaver. Tuberculosis was found in a dead animal. 

Introduction of Beavers into other parts of Germany, where they 
may find suitable living conditions and safety, is being considered. 
(Kriiger, 1931, pp. 53-56.) 

The present range in Germany is on the Elbe between Torgau 
and Magdeburg, and on the adjacent tributaries. Tuberculosis has 
been found the cause of death of a number of animals. Some Beavers 
have been introduced in the Schorfheide, near Berlin. (Reichsstelle 
fur Naturschutz, in litt., October, 1936.) 

In Wiirttemberg the last specimen was killed in 1869 on the 
Danube at the mouth of the Iller River (Wiirttembergische Natur- 
aliensammlung, in litt., October, 1936) . 

Blasius refers (1857, p. 407) to the former occurrence of Beavers 
in northwestern Germany on the Moselle, the Maas [now in the 


Netherlands], and the Weser; in the Luneburg area; and on the 
Schunter near Braunschweig. He also speaks of their more recent 
occurrence on the Havel and the Oder, in the Altmark, on the 
Vistula [now in Poland], in East Prussia, and at Schwenckfeld in 

"Harting says that 'at the close of the last century [eighteenth] 
many localities are reported to have been frequented by Beavers,' 
notably in Altmark, Preignitz, Middlemark, on the rivers Spree and 
Haxel and in the vicinities of Berlin, Potsdam, Oranienburg, Lieben- 
walde, Trebbin, Nauen, and Konigshorst. . . . Wagner in 1846 
mentions Beavers as living on the Danube, Amper, Isar, Iller, 
Salzach, and the Oder." (Millais, 1905, p. 161.) 

Denmark. The species was formerly distributed all over Den- 
mark, including Bornholm (Winge, 1908, p. 96) . 

Norway. Trade in Beaver skins was carried on early in the 
Middle Ages. Probably most of the Norwegian furs were exported 
to England. The species had begun to decrease by the close of the 
seventeenth century. In the middle of the eighteenth century it was 
probably still distributed throughout most of the woodland valleys, 
from the southernmost parts of the country to the farthest confines 
of Finmarken. 

In 1896 its range was chiefly in the Stifts of Christiania and 
Christiansand. The largest colony was located on the Nisser River 
in Nedenaes Amt. 

The trees felled are used both for food and for building material. 
The Beaver prefers the aspen (Populus tremula) and after that the 
birch, oak, and alder. 

According to an old superstition, the castor has the power to 
frighten away whales approaching a boat. In some parts the castor 
is worn from the garter as a specific for worms. In the sixteenth 
century the tail was regarded as a table delicacy. The teeth are 
worn as amulets in Finmarken, partly for ornament, partly as a 
protection against sickness, and were offered to the gods at the 
place of sacrifice and buried in the graves of heathen Lapps. 

By the middle of the nineteenth century the Beaver was fast 
becoming extinct, but the Game Laws of 1845 checked the decrease. 
By the end of the century it was on the increase and had extended 
its range by migration. The number was estimated at about 60 in 
1880, about 100 in 1883, and perhaps a few more in 1896. By this 
time Norwegian Beaver skins were very rarely in the market, and the 
castor was of little value. 

For a period of about 40 years after 1855 Beavers were allowed to 
be killed under certain restrictions. Then a closed season of 10 years 
was declared for the entire Amt of S0ndre Bergenhus and for the 
whole of Aamli. (Collett, 1898, pp. 105-126.) 


By 1931, as a result of state protection, the numbers on the Nidelf 
River had increased to 12,000 (Kriiger, 1931, p. 53). 

A limited amount of hunting is now allowed. During late years 
Beavers have been transplanted to the northern parts of the country, 
where they seem to thrive well. In some districts the farmers com- 
plain of damage to the forests by the Beavers. (Hj. Broch, in litt., 
December, 1936.) 

Sweden. The Beaver was formerly distributed all over the 
country, but gradually it became extinct. The last specimen in 
Smolandia was probably killed about 1800; that in Jemtland prob- 
ably about 1870. It was the value of the fur and the castor that 
caused its extermination. Beavers of the same race from Norway 
were first introduced in Jemtland in 1922, in Westerbotten in 1924, 
and in Wermland in 1925. These have all increased, and the number 
in Jemtland is now estimated at several hundred. There have also 
been introductions in other provinces, and further trials of this kind 
are planned. The animal is now very popular in Sweden, and its 
future appears to be rather promising, especially since it is protected 
throughout the year. (Einar Lonnberg, in litt., 1937.) 

Latvia. In former times the Beaver was found on many of the 
smaller rivers, but owing to excessive hunting it was exterminated 
about 1870. In 1927 the Government introduced four Norwegian 
Beavers in the State Forest of Kurland, and in 1936 two others in 
Smiltene in Livland. They have now increased in number to about 
40. Hunting is forbidden. (Forest Department of Latvia, in litt., 
March, 1937.) 

Lithuania. The species is probably exterminated. Since the 
World War two specimens have been illegally captured: one in 1921 
on the Dubisa, and one in 1935 on the Nemunas. Hunting is for- 
bidden. Reintroduction from neighboring countries is desirable. (T. 
Ivanauskas, in litt., November, 1936.) 

Poland. Game protective measures were instituted as early as 
the beginning of the eleventh century, when Boleslaus I the Great 
proclaimed an act for the protection of the Beaver (Benedyct Ful- 
inski, MS., 1933) . 

In ancient times and perhaps even in the seventeenth century 
the Beaver was quite common in all Poland. Owing to the reduction 
of forest areas and especially to the regulation of rivers, it retired 
to the eastern and northeastern parts of Poland, where it is still 
found. (M. Siedlecki, in litt., October, 1936.) 

Three preserves in the state forests, aggregating 684 hectares, are 
sanctuaries for Beavers. Another, the Bucharzewo Preserve, of about 
5 hectares, contains Canadian Beavers. (Benedyct Fulinski, MS., 


The present colonies of Castor fiber vistulanus all lie in the river 
basins of the Niemen, the Pripet, and the Dnieper, and perhaps also 
the western Duna. A map shows the present distribution at 14 
stations in the Niemen Basin and at 16 stations in the Pripet Basin. 
The last records on the middle course of the Vistula were in 1850; 
on the headwaters of the Vistula system, in 1861 ; on the headwaters 
of the Bug, in 1861 ; on the headwaters of the Dniester, in 1851. The 
largest colony in Volhynia in 1928 was estimated to contain 100 
individuals. In 1928-29 the total number in Poland was estimated 
at 235. The animals are now very strictly protected. (Kuntze, 1935, 
pp. 65-68.) 

Russia. Various early records are summarized by Nehring (1890, 
p. 105) as follows: Pallas (ca. 1770) reported that Cossack hunters 
sought Beavers on the steppe rivers of the Samara region, where the 
animals occurred very sparingly. According to Rytshkov, Beavers 
still existed in 1760-70 in the Bashkiri region; according to Evers- 
mann, about 1850 in the Perm Government; according to Kessler, 
at the same period in many rivers of the Kiev and Poltava Govern- 
ments; according to Krynitzki, about 1835 near Kherson on the 
lower Dnieper. 

Trouessart (1910, p. 130) includes northern Russia and southern 
Russia (Caucasus and rivers of the Caspian) in the range of the 
species. Millais states (1905, p. 161) that "Beavers were found on 
the Petchora and the Dwina in Russia until 1842, and possibly a few 
may still exist in their unfrequented tributary stream-." 

Of Russian Beavers we know comparatively little. In 1884, 566 
individuals were counted in the Rokitno Swamps. But by the time 
of World War I this number had greatly decreased, despite protec- 
tive measures instituted in 1911. It is doubtful if the colony set 
out on the Voronesh in 1886 still exists. (Kruger, 1931, p. 53.) 

The species was formerly widely distributed in the forested areas 
but is now almost exterminated and exists only in some reserves. 
These are in the Ukraine (on the Rivers Teterev, Soge, and Desna, 
on the tributaries of the Pripet, and in the former Government of 
Chernigov) ; in the Western Area and in White Russia ; and on the 
Usman in the former Government of Voronesh. In 1935 the total 
number of Beavers in the U. S. S. R. (including Siberia) was esti- 
mated at 2,500-3,000. (W. G. Heptner, in Hit., December, 1936.) 

Siberia. According to Eversmann (as reported by Nehring, 1890, 
p. 105), Beavers still existed about 1850 in the Baraba Steppe 
(between the Irtish and the Ob Rivers). "Gone from the Yenisei 
and Irtish, where formerly they were common, they were reported 
from the Pelyn, a tributary of the Obi, in Western Siberia, until 
1876, and they may still exist there" (Millais, 1905, p. 161). Troues- 
sart (1910, p. 130) includes Turkestan as well as Siberia in the 


range of the species. According to Millais (1905, p. 161), it was 
found even "as far east as Behring Straits." Kriiger states (1931, 
pp. 52-53) that it once ranged from the Urals to the Pacific Coast, 
but that the white examples with yellowish backs on the farther 
side of the Urals probably have not survived, and that there are 
reports of the complete extirpation of the species in Siberia. Ac- 
cording, however, to W. G. Heptner (in litt., December, 1936), it 
exists on the Rivers Konda and Sosva and their tributaries in the 
Ob Basin. 

""Schrenck (1859) reported the Beaver from Sakhalin, and he is 
quoted as authority by Aoki (1913, p. 298) and by Hatta (1928, 
p. 1036). The record is questioned, however, by Kuroda (1928, 
p. 224) , who calls attention to the lack of specimens. 

The paucity of beaver records from the Siberian wilderness sug- 
gests that the animal may never have been very abundant or thor- 
oughly distributed over that country. Perhaps the Siberian taiga, 
with its predominant coniferous growth, does not provide a sufficient 
quantity of the Beaver's favorite food trees, such as the aspen and 
other deciduous species, to support the animal in large numbers. 

Turkey, Syria, and Iraq. Danford and Alston (1880, p. 60) give 
an unsubstantiated report of a beaverlike animal in the marshes 
between Kaisariyeh and Indjesu, Turkey. They also cite reports by 
Heifer and Heifer (1879) of Beavers on the Euphrates near Aleppo, 
and by Schmarda (1853) in Mesopotamia. 

Persia. "The beaver, according to Eichwald, is common in the 
Araxes .... I insert it in the Persian fauna with some doubts." 
(Blanford, 1876, p. 51.) The above-mentioned report from the 
Araxes is categorically denied by later authors (Satunin, 1906, 
p. 374). 

Mongolia. In the upper Yenisei Basin, Tannu-Tuva, "a few 
beavers still exist in the upper tributaries of the Bei-Kem; but 
they are very rare, and their skins are seldom brought down to 
the markets. In old days they were mentioned as being included 
in a tribute sent by the Khan of the Ubsa region, then paramount 
chief of the Uriankhai tribes, to the Czar of Russia." (Carruthers, 
1913, p. 228.) 

The species "still exists . . . , it is said, in the highest tribu- 
taries of the Black Irtish in the Mongolian Altai" (Carruthers, 1913, 
pp. 630-631). 

Manchuria. "An animal recorded by Schrenck, but which does 
not appear to belong to the Manchurian fauna, is the beaver (Castor 
fiber) . It is true that skins of this animal have been secured from 
the natives in the Amur region, and that they find their way to the 
fur market in such places as Harbin and Mukden in Manchuria, but 
recent investigation tends to show that these skins have been brought 


from Alaska, having been bartered from one tribe of natives to 
another till they came into the hands of Russian or Chinese fur 
traders." (Sowerby, 1923, p. 170.) 

India. The Beaver may have occurred even in India, since, ac- 
cording to Buffon, the religion of the Magi forbade them to kill 
this animal (Blasius, 1857, p. 407) . 

Egypt. Since the species is supposed to be represented in the 
Egyptian hieroglyphs, it may have occurred in Africa (Blasius, 
1857, p. 407). 

Economics. "Had not the use of its hair in the manufacture of 
hats been superseded by that of silk, there is little doubt that the 
beaver, both in the Old World ^nd in America, would by this time 
have been numbered among extinct animals. As it is, the creature 
has but a hard time of it at best, for although there is no longer a 
demand for its hair by the hat-manufacturer, yet beaver-fur is an 
article highly valued by the furrier, and equally highly esteemed 
by the fair sex." (Lydekker, 1903, p. 244.) 

Family CRICETIDAE: Hamsterlike Rodents 

While various authors are not in accord on the limits of this family, 
it is probably safe to say that it consists of more than a hundred 
genera and more than a thousand forms. Representatives occur over 
the greater part of the world, and in general their numbers are legion. 
However, 14 New World forms are included in the preceding volume 
by Dr. Allen (1942) , and the 6 forms of the African genus Lophiomys 
are discussed here. A recent authority (Ellerman, 1941) makes a 
separate family (Lophiomyidae) of this genus. 

Genus Lophiomys Milne-Edwards: African Maned Rats 

The following remarks of the Committee of Experts (Hemming 
et al, 1938, p. 13), while naming only a single species, may be 
taken to apply to all known forms of this rare and peculiar genus: 

No species of rodent was included in either class of the Annex in the 
Convention of 1933, presumably owing to the small size and insignificant 
appearance of the majority of the species involved, and to the fact that they 
do not fall into the category of game animals. We see no reason however 
why a species of this Order should not be placed in the Annex if owing to 
their rarity they are in danger of extinction. 

A species of this Order which we should like to see protected is the Crested 
Bush Rat, Lophiomys imhausi Milne-Edwards, a very remarkable species 
living at altitudes of between seven and nine thousand feet in the mountains 
of Abyssinia and Kenya. This species which lives in pairs in dead trees and 
similar cover is peculiarly liable to attack and its numbers are known to have 
diminished considerably in recent years. 

We accordingly recommend that this species should be included in Class A 
of the Annex which, owing to the fact that rodents are not game animals, 


appears to us the most appropriate method of securing their protection. In 
the case of this particular species we shall hope to have received before the 
next meeting of the Conference the data to be collected by the Italian Scien- 
tific Mission. 

Hollister remarks (1919, p. 37) : "Although a few specimens of 
the maned rat find their way into collections from time to time, 
the animal is still so rare that no suitable series are available for 
study. If all the collections in various museums were combined it 
t would still be impossible to form any correct idea of the relation - 
'ships of the named forms, and it will doubtless be many years before 
sufficient material has accumulated." 

Under these circumstances the classification and nomenclature 
in the following accounts of the known forms of Lophiomys must be 
considered as no more than provisional. Possibly all the forms so 
far described will eventually prove to be no more than subspecifically 
distinct. All exhibit the same general color pattern. 

Sudan Maned Rat 


Phractomys aethiopicus Peters, Zeitschrift Gesammten Naturwissens. Halle, 
vol. 29, p. 195, 1867. (Based upon a skull from Maman, north of Kassala, 
Anglo-Egyptian Sudan.) 

FIG.: Anderson and de Winton, 1902, pi. 51. 

This species seems to be known chiefly from single specimens 
collected at no more than about half a dozen localities in the Anglo- 
Egyptian Sudan and Eritrea. 

Under the name of L. imhausi, Anderson .(in Anderson and de 
Winton, 1902, p. 289) describes an adult female from Erkoweet(?), 
on the mountains between Suakin and Sinkat, Sudan, somewhat as 
follows: Denser fur generally gray at the base, with a broad white 
band and wide brown tip; the long hairs broadly tipped with white; 
a triangular white area on top of the head, prolonged backward 
below the ears to the side of the neck, where the adpressed lateral 
band of yellowish hairs commences; a white spot below the eye; 
front and sides of head, throat, and sides of neck blackish brown; 
under surface generally pale brown, with an admixture of white; 
tip of tail white. Total length about 40 cm. (The brownish rather 
than blackish tone possibly represents a discoloration that had 
developed since the specimen was collected in 1880. A similar dis- 
coloration is now observable in the type of L. smithi Rhoads.) 

In writing of this specimen, Giglioli says (1881, p. 45) : "The 
Natives told Count Marazzani that the Lophiomys is rare, that it 
lives in deep holes in the strangely fissured rocks of that country." 
He also records a specimen killed at Keren in the Bogos country, 
Eritrea, in 1870. 


Oustalet (1902, p. 399) records a specimen from Massaua, 

"This animal is said to occur in the Khor Baraka and also at 
Tamai [respectively south and west of Suakin], and it is stated 
that it burrows under the roots of trees like a rat" (Anderson, in 
Anderson and de Winton, 1902, p. 290) . 

All, or nearly all, of the above-mentioned specimens, except the 
type of aethiopicus, were recorded as "L. imhausi" before the plas- 
ticity of the genus was recognized, but all of the localities are much 
nearer to the type locality of aethiopicus than to that of any other 
described form. 

Imhaus's Maned Rat; Imhaus's Crested Bush Rat 


Lophiomys Imhausii A. Milne-Edwards, L'Institut, vol. 35, p. 46, 1867. (Based 
upon a specimen secured alive at Aden, Arabia, but of unknown prove- 
nance (A. Milne-Edwards, 18676, p. 115) ; Thomas remarks (1910, p. 222) 
that Aden is "a place to which Somali animals are very commonly brought 
for sale.") 

SYNONYM: Lophiomys smithi Rhoads (1896) ("Sheikh Husein, West Somali- 
land" = Ethiopia) . 

FIGS.: A. Milne-Edwards, 18676, pis. 6-10; Kull, 1894, p. 136, fig.; Rhoads, 
1896, pi. 25; A. D. Smith, 1897, p. 64, fig.; Drake-Brockman, 1910, pi. 
facing p. 133. 

This species is " found probably throughout the Somali country, 
but [is] undoubtedly a very rare animal" (Drake-Brockman, 1910, 
p. 134). 

It is covered with very long silky hairs, of mixed white and black ; 
those of the back rising in a crest from the crown to the tip of the 
tail, and separated from those of the sides by an area of much shorter 
hairs, brittle and grayish tawny; tail long, not prehensile, covered 
with hairs like those of the body (A. Milne-Edwards, 1867a, pp. 46- 
47) . The general appearance is not ratlike. The dorsal crest is 
erectile. An adult male from British Somaliland measured: head 
and body, 11 inches; tail, 8 inches. (Drake-Brockman, 1910, p. 133.) 

The known distribution includes British Somaliland and south- 
eastern Ethiopia. 

Kull (1894) describes and figures two specimens from Somaliland. 

A specimen (the type of L. smithi) secured by A. D. Smith (1897, 
p. 64) at Sheikh Husein, Ethiopia, in 1894 was the only one seen in 
a journey of 4,000 miles through British Somaliland, Ethiopia, and 

In British Somaliland "I have seen it at Sheikh and near Burao, 
but never lower than 4,000 ft. One specimen was killed by Somalis 
at Upper Sheikh and one caught alive, while an adult female and 
young male were caught near Burao .... 


"Its custom of proceeding with crest erect is in all probability a 
protective measure to frighten its enemies, which might mistake 
it very easily for a young porcupine." (Drake-Brockman, 1910, 
p. 134.) 

Goba Maned Rat 


Lophiomys Bozasi Oustalet, Bull. Mus. Hist. Nat. [Paris], vol. 8, no. 6, 
p. 400, 1902. (Goba, southern Ethiopia; alt. 3,000 m.) 

This species appears to be known only from some three or four 

The female type is described as larger than L. imhausii; fur 
thicker; an elongate white spot over each eye, with a black band 
between; a white spot below the eye; muzzle and area about each 
eye black. Total length, 535 mm. (Oustalet, 1902, p. 401.) 

De Winton (in Anderson and de Winton, 1902, p. 291) records 
(under the name of L. imhausi) a specimen from near "Het Marafia" 
( = Let Marefia) and another from the forest of Tikem; both locali- 
ties are in Shoa, Ethiopia. 

Hollister (1919, p. 37) refers to the present species a specimen 
from Let Marefia, Shoa. 

Uaragess Maned Rat 


Lophiomys thomasi Heller, Smithsonian Misc. Coll., vol. 59, no. 16, p. 4, 1912. 
("Mt. Gargues (Uaragess), 6000 feet altitude, Mathews Range, British 

East Africa.") 

This species appears to be known from only three specimens from 
the type locality. 

"Allied most closely to ibeanus, differing chiefly in darker and 
more contrasting coloration .... General dorsal coloration deep 
black, the hairs everywhere broadly white tipped . . . ; the sides 
somewhat more extensively white than the median maned area; 
. . . lateral bands . . . olive-drab .... Head chiefly black with 
two prominent wide white bands over eyes, which meet on forehead, 
another large white spot below eye .... Underparts grayish, the 
hairs extensively white tipped . . . ; tail silvered like dorsal region, 
the extreme tip white. . . . Head and body, 270 mm., tail, 165." 
(Heller, 1912, p. 4.) 

"These three specimens were caught in rock crevices .... Heller 
believes these Lophiomys to be strictly rock-dwellers, notwithstand- 
ing reports of their living in holes of trees." (Hollister, 1919, p. 37.) 


Jackson's Maned Rat 


Lophiomys testudo Thomas, Ann. Mag. Nat. Hist., ser. 7, vol. 15, p. 80, 1905. 
("Ravine Station, [Mau Plateau,] British East Africa.") 

"The type-specimen . . . remains to this date unique" (Hollister, 
1919, p. 37). 

This species differs from the others in skull characters; "line of 
glandular bristles on sides narrower and less conspicuous than in 
other species" (Thomas, 1910, p. 223). "Basal third of underfur 
dark brown, middle third white, tip black, the contrast between the 
colours more marked than in L. Imhausi. Suborbital white spot well 
marked. Dark band dividing the frontal from the auricular white 
patch scarcely perceptible. . . . Hairs of lateral line olive. Under 
surface hoary grey .... Tail with its underfur mixed whitish and 
black, the tip for a length of about half an inch sharply contrasted 
white. . . . Head and body 296 mm.; tail 176." (Thomas, 1905, 
p. 81.) 

Mau Maned Rat 


Llophiomys] ibeanus Thomas, Ann. Mag. Nat. Hist., ser. 8, vol. 6, p. 223, 
1910. ("Mile 513 of the Uganda Railway (between Londiani and Lumbwa 
Stations) in Mau region," Kenya.) 

SYNONYM?: Lophiomys ibeanus hindei Thomas (1910). 

FIGS.: A. B. Baker, 1912, pi. 1; Hollister, 1919, pi. 2. 

This is perhaps the least rare species of Lophiomys. It occurs in 
the Mau region of Kenya, while the doubtfully distinct L. i. hindei 
has been recorded from the neighboring Aberdare Range and from 
Mount Kenya (Hollister, 1919, pp. 38-39) . 

This species "is coloured like the Abyssinian form referred to 
L. bozasi, and has equally prominent lateral stripes." It differs 
from other species in skull characters. (Thomas, 1910, pp. 223-224.) 

In addition to the type, specimens of L. i. ibeanus are recorded 
from El-Burgon and from the Mau Forest near Njoro, Kenya, while 
three specimens of L. i. hindei are recorded from the Aberdare 
Mountains (Thomas, 1910, p. 224). 

A. B. Baker (1912, p. 2) writes: 

This species of Lophiomys occurs in the higher part of British East Africa 
and is known only to the Wanderobo, a tribe of expert hunters, who explore 
every corner of the forests. Mr. Goldfinch was well acquainted both with the 
game of that region and with its animals generally, but this one he knew 
only from descriptions given by the natives. At his urgent request they secured 
two specimens in the forest near Nakuru, at about 8000 feet altitude. . . . 

Mr. Goldfinch states that Lophiomys is arboreal and lives in the thick 
forest of the high country, . . . also that the natives are averse to handling 


the animal, believing its bite to be poisonous. It is he says, "very rare or only 
got by accident here." ... It is strictly nocturnal. 

Lonnberg (1912, p. 100) records a specimen from Mau Escarp- 

Hollister (1919, p. 38) records specimens of ibeanus from the 
Naivashi Escarpment and from Nakuru, and one of hindei from 
Mount Kenya. 

"The first one of these animals I got was when I was stationed 
at Nakuru; it came from the Aberdare side. It was taken out of a 
hole in a tree by a Wanderobo .... I had no difficulty in getting 
all I wanted, and at one time I had something like a dozen of them." 
(Goldfinch, 1923, p. 1091.) 

Family MURIDAE: Old World Rats 

The limits of this family, as of the Cricetidae, are not definitely 
settled. The two families are similar in the multiplicity of their 
genera, species, and subspecies. While the Muridae were originally 
confined to the Old World, several forms of Rattus and Mus have 
attained world-wide distribution through transoceanic shipping and 
are thoroughgoing pests. In the genus Rattus, two species endemic 
on Christmas Island in the Indian Ocean have become extinct, and 
an Australian subspecies has apparently met the same fate. Single 
representatives of two other Australasian genera (Mastacomys and 
Zyzomys) are treated in the following pages. 

South Australian Spiny-haired Rat 


Rattus culmorum austrinus Thomas, Ann. Mag. Nat. Hist., ser. 9, vol. 8, 
p. 427, 1921. ("South Australia; type probably from Kangaroo Island." 
However, Iredale and Troughton (1934, p. 74) suggest "Port Lincoln" as 
the type locality.) 

More than a century has elapsed since the type specimen of this 
rat was sent to the Zoological Society of London in 1841, and 
there seem to be no records of appreciably later specimens. 

The fur is sparse, coarse, and more or less admixed with flattened 
spines; it is longer in this than in the other subspecies, the hairs of 
the back being commonly 20 mm. in length ; general color above gray 
rather than fawn color; under parts equally gray. Head and body, 
155 mm.; tail, 120 mm. (Thomas, 1921, p. 427; Jones, 1925, pp. 

Thomas (1921, p. 427) mentions six specimens besides the type, 
and remarks: "Evidently a common rat in South Australia in the 
forties, but whether it still exists in any out-of-the-way part of 
the colony we have no evidence to show." 


To this Jones adds (1925, p. 299) : "I know of no recent records 
or specimens of the species. So far, the out-of-the-way place has 
not been found by collectors, and this fact should prove a stimulus 
to our field naturalists." 

A. S. Le Souef writes (in Hit., February 15, 1937) that this par- 
ticular race is probably extinct, but that one or more of the other 
subspecies are still numerous at times. 

[The other subspecies are: Rattus culmorum culmorum Thomas 
and Dollman, of Queensland; R. c. youngi Thomas, of Moreton 
Island, Queensland; R. c. vallesius Thomas, of the interior of New 
South Wales.] 

Captain Maclear's Rat 


Mus macleari Thomas, Proc. Zool. Soc. London 1887, p. 513, 1887. (Christmas 

Island, eastern Indian Ocean.) 
FIGS.: Thomas, op. cit., pi. 42 (colored); Andrews, C. W., 1900, pi. 2 bis, 

figs. 1, 3, 6, 7, 8 (skull and teeth). 

This rat, isolated on Christmas Island, some 200 miles south of 
Java, the nearest land, is believed to have become extinct in the 
early years of this century. It is apparently nearest related to 
Rattus xanthourus of Celebes and R. everetti of the Philippines, 
which it somewhat resembles in appearance. 

About the size of a Roof Rat, it is described as grizzled rufous 
brown above, the belly but little lighter, pale rufous; longer hairs 
black, feet dark like the body. A striking feature is said to be the 
prominent long black hairs of the lower back, which, as in the other 
related rats, project far beyond the shorter portions of the pelage. 
The tail, which equals or slightly exceeds the length of head and 
body, is dark in its proximal half, white in its distal portion, and 
scaly. The skull is large and strongly built, with beaded supra- 
orbital edges, and the anterior edge of the zygomatic plate projects 
forward conspicuously. Measurements: head and body, 235-240 
mm.; tail, 246-267; hind foot, 48.5-50; ear, 17-17.5; basal length 
of skull, 47.5 ; zygomatic width, 26.2. Mammae four. 

This island rat was first made known by Thomas (1887) from a 
specimen brought from Christmas Island by Captain Maclear of 
the British surveying-ship Flying-fish, who procured it on his visit 
there in 1886. In the following year additional specimens were 
secured by J. J. Lister, who, as naturalist, accompanied a second 
expedition to the island on H. M. S. Egeria. At that time the island 
was uninhabited and covered with jungle and forest. Of about 40 
square miles in area, its highest point is about 1,200 feet above sea 
level; geologically, it is largely of coral limestone resting on a basis 


of volcanic rock. The specimens brought back by this expedition 
indicated a deposit of phosphate rock, to exploit which a settlement 
was founded at Flying-fish Cove, the only anchorage. Shortly after 
this, Dr. Andrews made a three-months' visit (in 1897) in order 
to survey the natural conditions there. His account contains prac- 
tically all that is known of the species, which was then by far the 
commonest of the mammals found in the island. He wrote: 

In every part I visited it occurred in swarms. During the day nothing 
is to be seen of it, but soon after sunset numbers may be seen running 
about in all directions, and the whole forest is filled with its peculiar 
querulous squeaking and the noise of frequent fights. These animals, like 
most of those found in the island, are almost completely devoid of fear, and 
in the bush if a lantern be held out they will approach to examine the new 
phenomenon. As may be imagined, they are a great nuisance, entering the 
tents or shelters, running over the sleepers, and upsetting everything in their 
search for food. They seem to eat anything, and destroy any boots or skins 
incautiously left within their reach. Their natural food appears to be mainly 
fruits and young shoots, and to obtain the former they ascend trees to a 
great height. ... In the settlement they utterly destroy all the fruit they 
can get at, and frequently come into conflict with the fruit-bats on the 
tops of the papaia-trees. A number of dogs is kept to keep them in check, 
and near the settlement they are certainly already less numerous than else- 
where. In the daytime these rats live in holes among 1 the roots of trees, in 
decaying logs, and shallow burrows. They seem to breed all the year round. 

After 10 years' absence, Andrews (1909) again visited Christmas 
Island for the purpose of ascertaining what changes had taken 
place in the interim as a result of white occupation. Such changes 
were "chiefly noticeable in the immediate neighbourhood of the 
settlement and quarries, while the rest of the island, although tra- 
versed by roads in several directions, is practically unchanged." The 
rats, however, had gone. For whereas 10 years earlier they were 
found everywhere all over the island in abundance, in 1908, in spite 
of continual search, not a single specimen of this tree-climbing 
species or of the other burrowing rat, R. nativitatis, could be found 
in any part of the island. He says further: 

This complete disappearance of two such common animals seems to have 
taken place within the last five or six years, and to have been the result of some 
epidemic disease, possibly caused by a trypanosome, introduced by thfe 
ship-rats. These are a variety of Mus rattus, and have been introduced in 
considerable numbers, though they do not seem to have spread to the 
remoter parts of the island at present, at least to any great extent. The 
disappearance therefore of the native forms cannot be due to direct com- 
petition with the intruders, but must be the result of disease, a conclusion 
supported by an observation made by the medical officer, Dr. McDougal, 
who told me that some five or six years ago he frequently saw individuals 
of the native species of rats crawling about the paths in the daytime, 
apparently in a dying condition. 

Since Andrews's second visit in 1908, one or two other zoologists 
have visited Christmas Island for the study of its fauna, notably 


M. W. F. Tweedie in 1932, but apparently no one has since found 
a trace of the two indigenous rats or of the shrew (Crocidura 
fuliginosa trichura) that were abundant before the settlement. No 
doubt the supposition that they were exterminated through the 
spread of some disease brought in by introduced House Rats is the 
most likely explanation of their disappearance. Chasen (1933) , who 
has written of the birds of the island, adds that in addition to Rattus 
rattus (subsp. ?) , the House Mouse (Mus musculus) and the small 
Rattus concolor, a member of a group adaptable to colonization as a 
human acolyte, have also been introduced in this island. 

G. M. A. 

Christmas Island Burrowing Rat; "Bulldog Rat" 


Mus nativitatis Thomas, Proc. Zool. Soc. London 1888, p. 533, 1889. (Christ- 
mas Island, eastern Indian Ocean.) 

FIGS.: C. W. Andrews, 1900, pi. 2 (col. fig.) ; pi. 2 bis, figs. 2, 4, 5, 9, 10 (skull 
and teeth). 

This rat and Rattus macleari (q. v.) are the only indigenous ter- 
restrial rodents known from Christmas Island, Indian Ocean, and 
are both now believed to be extinct. 

Rattus nativitatis was first collected by J. J. Lister in 1887, when 
as naturalist aboard H. M. S. Egeria, in the year following the 
visit of Captain Maclear, he explored part of the island. On this 
occasion, a landing party under Captain Aldrich cut a way through 
the jungle to the highest part of the island. 

In contrast to the other species, R. macleari, this rat was a more 
stoutly built animal of burrowing habits. It is described as a large 
species about 17 inches in total length with a tail much shorter than 
head and body, of a thickset clumsy form, but having a peculiarly 
small and delicate head. In color it was a dark umber brown all 
over, the belly not or scarcely paler. The fur of the back, though 
long, thick, and coarse, was without the elongated piles characteristic 
of R. macleari. The claws were broad and strong, adapted for 
digging. Mammae abdominal, three pairs. There is a slight degree 
of variation in color, some individuals being a warmer brown than 
others, and occasional ones having a small irregular patch of white 
fur on the belly. Teeth relatively small and weak. Measurements: 
head and body, 275 mm.; tail, 182; hind foot, 50; ear, 24 (these 
for the largest of nine specimens). Skull: basal length, 46.8 mm.; 
zygomatic width, 24.8; nasals, 20.5; diastema, 15.5; upper cheek 
teeth, 7.6. A comparison of the forearm and hand bones in the two 
species is given by Forsyth Major (with figures) in Andrews's 
(1900) Monograph of Christmas Island. 


This rat apparently was less generally distributed over the island 
than R. macleari, inhabiting hilly areas in the interior. Andrews 's 
account supplies practically all that is known of it. He wrote: 

Though very numerous in places, especially on the hills, e. g. Phosphate 
Hill, [it] is very much less common than M. macleari. I never saw one in 
Flying Fish Cove [the settlement], though they certainly have been killed 
there. They seem to live in small colonies in burrows, often among the 
roots of a tree, and occasionally several may be found living in the long, 
hollow trunk of a fallen and half-decayed sago-palm (Arenga listeri). The 
food consists of wild fruits, young shoots, and, I believe, the bark of some 
trees. [It is a] much more sluggish animal than M. macleari, and unlike it, 
never climbs trees; and it is difficult to avoid the belief that the former 
species is being supplanted by the latter in spite of the abundance of food. 
Both animals are strictly nocturnal, and M. nativitatis, when exposed to 
bright daylight, seems to be in a half-dazed condition. The Ross family in 
Christmas Island have given this species the name "Bull-dog Rat," and this 
has been adopted by the Malays. 

This was in 1897. When, in 1908, Andrews revisited the island 
to see what changes had followed the planting of a settlement there, 
he found both species apparently quite gone. "In spite of continual 
search, not a single specimen of either species could be found in any 
part of the island." This disappearance, as detailed under Rattus 
macleari, was conjectured to have taken place about five or six years 
earlier, when the medical officer stationed there had frequently seen 
individuals of the native rats "crawling about the paths in the day- 
time, apparently in a dying condition." Andrews suggests that the 
introduced Roof Rat, by then already present in considerable num- 
bers, had brought in some epizootic disease to which the native 
species had been susceptible, and in consequence they had been 
entirely wiped out in the brief space of a few years. (Andrews, 1909, 
pp. 101-102.) 

At the time of Andrews's first visit he wrote (1900) : "The con- 
ditions of life are apparently extremely favourable, food being 
always abundant, and the hawk and owl, which are the only possible 
enemies [of these rats], feeding mainly on birds and insects. The 
consequence of this is that all the species of mammals are extremely 
common, and the individuals are always exceedingly fat. Perhaps 
Mus [= Rattus] nativitatis, the bull-dog rat as the Cocos Islanders 
have named it, is the least numerous, probably because of some 
competition with the much more active and versatile M. macleari, 
but most specimens of M. nativitatis have a layer of fat from half 
to three-quarters of an inch thick over most of the dorsal surface 
of the body." Possibly this very abundance of individuals and their 
fat condition made them the more susceptible to any disease brought 
in from outside. 

While conjecture as to the origin of the endemic fauna is more or 
less futile, Andrews nevertheless points out that on the whole its 


relations are with "Austro-Malayan" rather than with Javan types. 
He noticed on several occasions the transport of insects to the island 
by storms "which, during the rainy season, blow occasionally from 
the northern quarter," but inclines to the supposition that the "rats, 
the fruit-bat, and possibly some of the land birds, very probably 
owe their introduction to the island" to the transport by rafts of 
trees brought by the equatorial drift from the Timor Sea. This 
island is at least of unusual interest as affording a case in which 
the native fauna has within a few years been altered as a result of 
settlement by man, and two of its few native mammals have be- 
come extirpated. 

G. M. A. 

Broad-toothed Rat 


Mastacomys juscus Thomas, Ann. Mag. Nat. Hist., ser. 5, vol. 9, p. 413, 
1882. ("Tasmania.") 

This rare rat survives in the Otway Forest of Victoria and in 
Cradle Valley, Tasmania, but little or nothing is known of its present 
occurrence elsewhere. 

Fur long and soft; general color dark grayish brown above and 
below, the dorsal hairs tipped with light brown and the ventral 
hairs with white; tail and feet dark brown. Head and body, 142 
mm.; tail, 95 mm. (Thomas, 1882, pp. 413-415.) It is a large rat, 
with a stout build, strong limbs, and a short tail. Head and body 
(Tasmanian specimens), 170-182 mm.; tail, 110-113 mm. (Finlay- 
son, 1933a, pp. 126, 128.) Victorian specimens have longer tails 
(119-124 mm.) (Brazenor, 1934, p. 161). 

The type specimen, from an unspecified locality in Tasmania, was 
acquired by the British Museum in 1852, and for 80 years no further 
information seemed to be forthcoming as to its occurrence in that 
state. In 1931, however, five specimens were collected in Cradle 
Valley, northwestern Tasmania, at an altitude of about 3,000 feet. 
The Broad-toothed Rats were living in colonies in grassy areas on 
open heaths, in association with Eastern Swamp Rats (Rattus 
lutreolus). "Both rats are probably quite numerous, but the laby- 
rinths are the chosen hunting grounds of Dasyurus vivverinus [sic], 
and it was not until several days trapping had got rid of the latter 
that rats began to be caught." (Finlayson, 1933a, pp. 125-126.) 
Doubtless this Native Cat acts as a check upon the increase of 
the rats. 

Lydekker (1885, p. 227) records some bone fragments of Masta- 
comys juscus from the caves of the Wellington Valley, New South 


Brazenor (1934, pp. 159-160) records specimens from the follow- 
ing localities in Victoria: Swan Island; Gippsland; Layer's Hill 
in the Otway Forest; Olangolah, near Beech Forest, at the head 
of the Gellibrand River. He also mentions a specimen, apparently 
previously overlooked, from the "West Coast of Tasmania, 1872." 
He writes (in litt., March 3, 1937) that the species "still survives 
in the Otway Forest but not in any numbers." 

While-tailed Rat 


Mus argwrus Thomas, Ann. Mag. Nat. Hist., ser. 6, vol. 3, p. 433, 1889. 
("South Australia.") 

This South Australian rat is apparently known only from the 
type specimen. 

The fur is crisp; general color above pale sandy rufous; ears 
rounded, thinly covered with fine white hairs; muzzle and under 
parts white, the line of demarcation on the sides not sharply defined ; 
hands and feet pure white ; tail uniform white above and below, the 
tip slightly penciled. Head and body, 83 mm.; tail, 101. (Thomas, 
1889, pp. 433-435.) 

"Of this very distinct species there is no material available in 
South Australia, which was the home of the type specimen. . . . 

"There seem to be no recent records of this remarkable little 
rat, and no observations on its habits. Probably it is one of the 
many lost species of which no specimens are preserved in our State 
collections." (Jones, 1925, pp. 336-337.) 

A. S. Le Souef remarks (in litt., February 15, 1937) that these 
native rats do not stand up to settlement or invasion of their habitat 
by Rattus rattus. 

E. Le G. Troughton writes (in litt., April 16, 1937) that the lack 
of records since the original description in 1889 supports Wood 
Jones's conclusion that the species may be lost from the state. 

[According to L. Glauert (in litt., March 17, 1937) , the Western 
Australian subspecies, Z. a. indutus (Thomas), is "not reduced in 

Order CARNIVORA: Carnivores 

Family CANIDAE: Wolves and Foxes 

The Canidae are nearly cosmopolitan, indigenous species being 
found in all important land masses except Madagascar, the central 
and eastern parts of the Malay Archipelago, New Zealand, and 


Tasmania. There are about 19 'genera and more than 200 species and 
subspecies. The predatory habits of some of the larger species 
(especially the wolves) bring them into conflict with the economic 
interests of man, and the animals have suffered accordingly par- 
ticularly in North America. Accounts of no less than 24 New World 
forms appear in Dr. Allen's volume (1942), while only 4 Old World 
forms are treated herein. One of the these, the Japanese Wolf, is 

Abyssinian Wolf; Abyssinian Red Wolf. Cuberow (Ethiopian) 


Canis simensis Ruppell, Neue Wirbelthiere zu der Fauna von Abyssinien 
gehorig, Saugethiere, p. 39, 1835. (Mountains of Simien, Abyssinia.) 

FIGS.: Ruppell, 1835, pi. 14; Mivart, 1890, pi. 6; Bryden, 1899, pi. 15, fig. 8; 
Lydekker, 1908, pi. 15, fig. 8; Fuertes, Abyssinian Birds and Mammals, 
pi. 29, 1930. 

This interesting animal has a restricted range and occurs in 
limited numbers. The Committee of Experts of the Second Inter- 
national Conference, held at London in 1938, states (1938, p. 8) that 
this species is "almost completely confined to Abyssinia," and sug- 
gests its inclusion in Class A of the Annex at the next Conference 
for the Protection of African Fauna and Flora. 

Snout long and slender; general color light yellowish reddish 
brown, mixed with black on the sides; white about the mouth, eyes, 
inner margins of ears, chest, belly, lower parts of limbs, and lower 
side of tail toward base ; distal half of tail blackish, and upper side 
toward base mixed with black. Head and body, 99 cm. ; tail, 25 cm. 
(Mivart, 1890, pp. 18-19.) 

"We observed this wolflike dog in the mountains of Simien, 
where it lives in packs, and hunts tame sheep and small game, but 
never becomes dangerous to man. It occurs also in most of the 
other Abyssinian provinces. Its vernacular name in Simien was 
given to me as 'Kaberu.' ' (Ruppell, 1835, p. 39, transl.) 

"Since Riippell's time little has been heard of this wolf and scarcely 
any fresh or recent information is to be obtained concerning it. From 
its predatory habits it is probable that the Abyssinians, so soon as 
they began to acquire fire-arms, turned their attention to its destruc- 
tion, and that in consequence it has become much scarcer than it 
used to be. ... 

"It would be extremely interesting to know if this handsome wolf 
still survives -in Abyssinia in any numbers. Modern travellers and 
sportsmen apparently make no mention of it." (Bryden, 1899, 
pp. 601-602.) 

Lydekker (1908, p. 462) refers to "its rarity and zoological in- 


terest." The Cuberow "was scarcely known in England, except by 
its skull, till a few years ago, when skins were brought home by 
Major Powell-Cotton." The latter saw several of the animals alive 
in the mountains of Simien. 

Maydon (1932, pp. 220-221) writes that "the Red Wolf is com- 
mon" at Simien, and refers to it as being seen occasionally on the 
Gojam plateau, between Lake Tsana and Addis Ababa. 

W. H. Osgood (oral communication, 1936) speaks of this species 
as not uncommon locally. The Field Museum expedition of 1926-27 
obtained about five specimens. Alfred M. Bailey (oral communi- 
cation, 1937) does not consider that it is in any danger from the 

This animal appears to occupy a peculiar zoological position. 
Lydekker considers it neither a wolf nor a jackal, while Pocock 
denies to it affinity with the foxes. 

[A subspecies from south-central Ethiopia has been proposed by 
De Beaux under the name of Cam's (Simenia) simensis citernii (Atti 
Soc. Ital. Sci. Nat., vol. 61, p. 25, 1922; type locality, "Arussi: 
Barofa"). No information is at hand concerning its numerical 

Japanese Wolf 


Canis hodophilax Temminck, Tijdschr. Natuurl. Geschied. Physiol., pt. 5, p. 284, 
1839. (Japan; i. e., Hondo.) (Cj. Harper, 1940, p. 192.) 

FIGS.: Temminck, 1842-45, pi. 9; Mivart, 1890, p. 14, fig. 17; Beddard, 1902, 
p. 418, fig. 209. 

This wolf is now considered extinct. 

It is distinguished from the European Wolf by its smaller size 
and shorter legs, though it differs but little in the nature and color 
of its pelage; fur short and smooth, but tail bushy; ground color 
gray or ashy; basal two-thirds of the hairs of back and rump thus 
colored, the tips black ; sides, neck, belly, and tail gray, the extreme 
tips of the hairs blackish; head and muzzle dark gray; lips more 
or less whitish ; outer surface of ears brownish rufous ; four extremi- 
ties gray, washed with rufous and brown; tail tip without colored 
tuft. Height at shoulder, 16 inches; total length, 3 feet 9 inches, of 
which the tail comprises about 1 foot; ears, 3 inches. (Temminck, 
1844, pp. 38-39.) "Prof. Brauns . . . says that in the Museum at 
Tokio there are very differently coloured skins, namely 'yellowish/ 
'brownish,' and 'whitish grey' " (Mivart, 1890, pp. 14-15). 

The Japanese Wolf lives in wooded and mountainous regions, and 
hunts in small family parties. It is as much dreaded by the Japanese 


as the European Wolf is in its range. It shows itself often in winter, 
notwithstanding the assiduous pursuit of which it is the object. The 
Japanese state that its flesh is unwholesome. (Temminck, 1844, 
p. 39.) 

A female wolf from Japan was presented to the Zoological 
Society of London in 1878 (Flower, 1929, p. 114) . Mivart (1890, 
p. 15) records a skull in the British Museum from the province of 

Thomas (1906, p. 342) records a specimen collected in 1904 or 
1905 in the vicinity of Washikaguchi, Nara Ken, Hondo. The col- 
lector, M. P. Anderson, adds: "The Wolf was purchased in the flesh, 
and I can learn but little about it. It is rare, some say almost 

Aoki (1913, p. 317) gives the range of this animal as "Hondo 
(Thomas), China." Hatta remarks (1928, p. 1033): "Cam's hodo- 
phylax T. confined in Japan to the heart of Hondo, Yamato and 
Wakayama, occurs also in China." These reports from China are 
considered erroneous. Pocock (1935, p. 658) records a skull from 

Nagamichi Kuroda writes (in litt., July 5, 1938) that many of 
these wolves were formerly said to be seen in the mountainous dis- 
tricts of Hondo, but that the animal is now considered completely 
extinct. It was destroyed because of its injuriousness to men and 
cattle. It is said that the only specimens in Japan are a mounted 
male from Fukushima Prefecture, Hondo, which is now preserved 
in the Tokyo Science Museum, and one or two skulls. 

Kuroda (1938, p. 36) records the animal from the following addi- 
tional localities in Hondo: Rikuchu, Shimotsuke, and Aomori. 

Yezo Wolf 


Cam's lupus hattai Kishida, Lansania, vol. 3, no. 25, p. 73, 1931. (Sapporo, 

Hokkaido, Japan.) 
SYNONYM: Cants lupus rex Pocock (1935). 

Although extinct in Hokkaido (or Yezo), this wolf survives in 
Sakhalin and perhaps in the Kuriles. 

It is much larger than Cam's hodophilax of Japan and is dis- 
tinguished from C. I. lupus of Europe by its larger premolar teeth 
and by its longer palate and mandible (Pocock, 19356, p. 659). 

In the Amur region, according to Schrenck (1859, pp. 45-48), 
the wolf is most numerous in northern Sakhalin. Its principal object 
of chase is the wild Reindeer. Occasionally packs approach the 
villages or solitary houses of the natives and destroy their dogs. The 
animal ranges to the south end of Sakhalin and occurs also on the 


Pocock (1935b, pp. 659-660) refers to "the discovery of this big 
wolf in Yeso in the early 'eighties/' and to the opinion of Brauns 
(1881) "that it possibly inhabited the Japanese islands . . . between 
Yeso and Kamschatka." 

"Canis lupus L. is found ... in Sakhalin; in Hokkaido it was 
abundant some thirty years ago, but it has decreased so that it seems 
to be totally exterminated at present" (Hatta, 1928, p. 1037) . 

"Aoki and Kishida both reported it from this island [Sakhalin] 
and Hokkaido (rare) " (Kuroda, 1928, p. 226) . 

"In authentic historic times the wolves occurred in the main 
island of Hokkaido, in Sakhalin, and in Kunashiri, Etoruf and 
Paramushir of the Kurile Islands. It seems true that the wolves 
were not so frequent in Hokkaido as compared with the other 
mammals. They were still fewer in Sakhalin and in the Kurile 
group. Though old records say that the wolves fed mostly upon the 
deer which abounded in Hokkaido, at the beginning of settlement 
they wrought serious havoc amongst herds. So the government at 
that time paid a high bounty for the slaughter of the animal. For 
instance, the local government in Sapporo paid 7 yen for one wolf 
from 1878 to 1882 and 10 yen for each from 1883 to 1885. More 
than 1500 wolves were brought in for the bounty during the 11 years 
from 1878 until 1888. Since then we have heard scarcely any account 
of the animal in Hokkaido." (Inukai, 19326, p. 525.) 

Kuroda (1938, p. 36) gives the range of this subspecies as Sak- 
halin, the Kuriles, and Hokkaido ; on this last island it is extinct. 

Japanese Raccoon-dog 


Canis viverrinus Temminck, Tijdschr. Natuurl. Geschied. Physiol., pt. 5, 

p. 285, 1839. (Japan.) 
FIGS.: Temminck, 1842-45, pi. 8; Martens, 1876, pi. 1. 

Formerly abundant in Japan, this animal has become extremely 

The form is small and foxlike; the tail is short and bushy. The 
general color is yellowish brown; hairs of the back, shoulder, and 
tail tipped with black; arms and legs blackish brown; a large dark 
brown spot on each side of the face, beneath and behind the eye 
(Martens, 1876, p. 78). The measurements of some representative 
of the species on the Asiatic mainland are given by Mivart (1890, 
p. 135) as follows: head and body, 530 mm.; tail, 140 mm. 

A century ago the Raccoon-dog was considered very common in 
Japan (Temminck, 1844, p. 40) . At this period "Siebold found it to 
be very common throughout the Japanese islands, where its flesh 
was considered as good food with an agreeable flavour, and its 


powdered, calcined bones a valuable medicine. ... It is not deemed 
destructive to poultry. The natives employ its skin to make bellows, 
and also to decorate their drums and for winter head-gear." (Mi- 
vart, 1890, p. 135.) 

Possibly Ognev refers to its former rather than to its present 
status when he writes (1931, p. 369) that it is widely distributed in 
Japan and particularly common on Hondo (Honshiu) Island. 

In the open season extending from October 15, 1929, to April 15, 
1930, 15,218 of these animals were taken in Japan (Uchida, 1935, 
p. 8.) 

FIG. 22. Japanese Raccoon-dog (Nyctereutes procyonoides viverrinus} 

"Muko-jima, a small island in the Inland Sea on the coast of 
Yamaguchi Prefecture, is famous as a sanctuary for the raccoon dog 
.... In feudal times it was abundant throughout this country. 
Owing to indiscriminate hunting it became extremely scarce." (Ka- 
buraki, 1934, pp. 4183-4184.) 

Uchida writes (1935, p. 25) that the animal is gradually becoming 
scarce in Japan, since its fur is highly valued and a large number 
of skins are exported annually. Mukojima is inhabited by innumer- 
able individuals. It is supposed that persecution on the Japanese 
mainland gradually forced them to migrate to the island, where 
they found a safe breeding place and an abundance of fish for food. 

[Concerning the several mainland representatives of this species, 
Arthur de C. Sowerby writes (in litt., April 24, 1937) : 

"There are now several subspecies of the raccoon-dog recognized in 
China and neighbouring regions, namely, Nyctereutes procyonoides 


procyonoides (Gray) of Central and South China, Nyctereutes pro- 
cyonoides orestes Thomas of South-west China, and Nyctereutes 
procyonoides ussuriensis Matschie of North-east China, Manchuria, 
and neighbouring eastern Siberia and Korea. Except in a few places 
these animals are probably as plentiful to-day as they ever were, 
except that they seem to have been badly decimated in the Amur 
region, where once they were particularly plentiful, for the sake of 
their pelts. In years past large quantities of the latter have been 
exported from China, but at present there is no demand on either the 
American or European markets for the 'raccoon' or 'raccoon-dog' 
skins, as they are known to the trade, and consequently none are 
coming from the interior to the ports, except such as are required 
for the home market, which is fairly considerable." 

A few years ago approximately one-half million skins were ex- 
ported annually from Shanghai (Sowerby, 1934a, p. 287).] 

Family URSIDAE: Bears 

The bears occur on all continents except Australia and perhaps 
Africa (where a single problematical, extinct species has been re- 
ported from Morocco and Algeria). The only South American 
species is restricted to the Andean region. Thus the distribution of 
the Ursidae is not quite so extensive as that of the Canidae. The 
bears have suffered perhaps even more than the wolves at the hands 
of man. Seven genera and about 135 forms are provisionally recog- 
nized. The majority of the latter, however, are North American 
Grizzly Bears, whose exact taxonomic status remains somewhat 
uncertain. Dr. Glover M. Allen, in his volume on New World mam- 
mals (1942), treats all the Grizzly Bears and some of the Black 
Bears, as well as the South American Spectacled Bear. The present 
volume deals with the various forms of the Brown Bear in Europe 
and Asia and with the Atlas Bear of North Africa. 

Old World Brown Bear. Ours brun (Fr.) . Brauner Bar (Ger.) . 
Oso (Sp.). Orso bruno (It.) 


[Ursus] arctos Linnaeus, Syst. Nat., ed. 10* vol. 1, p. 47, 1758. (Sweden.) 
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mamm., vol. 5, pi. 211 (U. a. pyrenai- 
cus), pi. 212 (U. a. collaris), 1824; Gervais, Hist. Nat. Mammif., pt. 2, 
pi. 13, 1855 (U. a. pyrenaicus) ; Fitzinger, Bild.- Atlas, Saugth., fig. 72, 
1860; Millais, 1904, pi. facing p. 236; Martin, 1910, pi. 38; Cabrera, 1914, 
pi. 5 (U. a. pyrenaicus} ; Ognev, 1931, pi. 1 (U. a. caucasicus) ; Pocock, 
1932, pi. 2, upper fig.; Castelli, 1935, pis. 1, 2. 

Numerous forms of the Brown Bear have been described, but 
there is no general agreement on the validity of most of them, and 


the systematics of the group remains in a state of very considerable 
confusion (cf. Miller, 1912, pp. 285, 296; Ognev, 1931, pp. 14-118; 
Pocock, 1932). In the account that follows, only two of the forms 
will call for separate treatment. Among those that have been de- 
scribed, the following may be mentioned as being better known or 
as having gained more or less recognition in the literature: 

Ursus arctos arctos Linnaeus, of northern Europe. 

U. a. pyrenaicus Fischer, of the Pyrenees and northern Spain. 

U. a. meridionalis Middendorff, of the Caucasus. 

U. a. syriacus Hemprich and Ehrenberg, of Asia Minor. 

U. a. collaris Cuvier, of Siberia. 

U. a. beringianus Middendorff, ranging from Manchuria to Kam- 

U. a. lasiotus Gray, of Mongolia, Manchuria, Hokkaido, and the 

U. a. isabellinus Horsfield, of the western Himalayas and the 
Thian Shan. 

U. a. pruinosus Blyth, of Tibet (not generally regarded as con- 
specific with U. arctos, but so treated by Pocock, 1932). 

The Brown Bear has become extinct over the greater part of its 
former range in western Europe but survives in small numbers in 
remote and chiefly mountainous areas in Norway, Sweden, Spain, 
France, Austria, Czechoslovakia, Italy, Yugoslavia, Greece, Bul- 
garia, Rumania, Poland, and Estonia. It has remained much com- 
moner in parts of Russia and northern Asia. 

The general form of the species is short and heavy; fur long and 
rather loose; head moderately pointed, broad posteriorly; ear short 
and rounded; front claws strongly curved, blunt, at least twice as 
long as hind claws ; tail very short, concealed in the fur. The general 
color is usually a light brown or dull buff, the head not essentially 
different, but feet and outer surface of legs darker. There are many 
individual and racial variations in color. Measurements of an adult 
male from Sweden: head and body, 1,900 mm.; tail, 80 mm.; hind 
foot, 195 mm.; ear, 90 mm. (Miller, 1912, pp. 287-296.) 

The range of the species is the "entire continent of Europe wher- 
ever sufficiently extensive forests remain; east into Asia" (Miller, 
1912, p. 287). 

Great Britain. Numerous postglacial remains have been found 
in various parts of England. These include bones from refuse heaps 
that are probably of Roman origin. The remains found in Ireland 
appear to belong to an older species than Ursiis arctos. In ancient 
times in Britain the animal was trailed with boar-hounds and at- 
tacked with arrows, pikes, clubs, javelins, and long knives. The 
great Caledonian forest in Scotland seems to have been the chief 


stronghold of the British bears. Bears were transported from Britain 
to Rome, probably in the fourth century. After the extinction of 
the species in Britain, foreign animals were imported for the pur- 
pose of "bear-baiting." This was done in the reigns of Henry II, 
Henry VIII, Queen Elizabeth, and Queen Anne. Queen Elizabeth 
entertained the French Ambassador and the Danish Ambassador 
at different times with such a spectacle. (Harting, 1880, pp. 11-29.) 

Many place names in Wales afford evidence of the former occur- 
rence of bears. Boyd Dawkins thinks that they became extinct in 
Britain before the tenth century. Bell, in his British Quadrupeds, 
says that they may have existed in Scotland as late as 1073, but 
tradition gives the latest date as 1057. (Millais, 1904, pp. 236-237.) 

Norway. According to Collett, the bear was numerous about 
1750 and did great damage to the cattle all over the country, but 
during the following 150 years the numbers were reduced owing to 
improved firearms and to high rewards paid for animals shot (Hj. 
Broch, in litt., December, 1936). 

Bowden states (1869, p. 4) that "the Brown Bear ... is pretty 
common in all parts of this country, especially in Nordland and the 
central districts." 

"The average number that is killed yearly amounts to ... 250" 
(Barnard, 1871, p. 262). 

"The number of bears in Norway is now beyond doubt very small. 
... A conservative estimate would be that there are not more 
than 20-25 bears in Norway south of the Trondhjem Fjord. In the 
northern parts the number is beyond doubt also very small, chiefly 
consisting of stray bears from Sweden and Finland. . . . 

"We have done everything in our power to attempt a protection 
sufficient to stop the extinction which obviously threatens the species 
in this country. But all attempts have been without avail, as it 
cannot be denied that the bears occasionally do some damage to 
sheep .... 

"We have attempted the establishment of a reservation in certain 
forest tracts owned partly by private owners and partly by the 
government. But the project has until now failed." (Harald Platou, 
in Hit., November 22, 1932.) 

By a law of 1932 the bear may be killed only from May 15 
to November 1. Previously "the bear, being considered a pest, re- 
ceived no protection at all. In fact there was a premium for killing 
him .... 

"This recent protection ... is due to the efforts of the Norwegian 
Association for Hunting and Fishing, which endeavored to induce 
the Storting to protect the bear all the year round. The Association 
had even collected money which it offered to place at the disposal 
of the Norwegian Department of Agriculture to compensate for 


losses to farmers and goat and sheep owners for damage done by 
the bear. . . . This offer was not accepted. . . . 

"It is not known exactly how many bears there are in Norway. 
. . . The greatest number are to be found in the tract Eastern 
Hallingdal the Hemsedal mountains to Laerdal, where it is esti- 
mated that there are from 15 to 25 bears. It is possible that there 
are a few bears between Valdres and Gudbrandsal and in the central 
part of Telemarken, but their number is uncertain. There are also 
a few bears in the northern part of Norway which come from Swe- 
den." (Julius Wadsworth, in litt., July 28, 1932.) 

"An area of about 125,000 acres north of the city of Lillehammer 
has been privately set aside for the protection of the bear." Mr. 
Platou "thinks the bears will now be preserved from extinction." 
(Julius Wadsworth, in litt., May 9, 1933, and July 20, 1933) . 

Sweden. The bear was formerly found in all parts of the coun- 
try, but in most of the provinces constituting Gotaland it had prac- 
tically disappeared during the eighteenth century. ^ It has been 
calculated that in the whole country 1,351 bears were killed during 
the period 1827-1836, and 1,055 during the period 1847-1856. In 
the 50-year period from 1856 to 1905, 2,762 bears were killed in 
Sweden, including 86 in Wermland, 499 in Dalecarlia, 171 in Gavle- 
borg, 144 in Westernorrland, 796 in Jemtland, 292 in Westerbotten, 
and 770 in Norrbotten. The rapid decrease is illustrated by the 
following statistics on the numbers killed in six of the above- 
mentioned districts: 908 in 1856-1865; 434 in 1876-1885; 109 in 
1896-1905. In olden times a small bounty was paid on each bear 
killed, and in 1864 the amount was raised to 50 riksd. Sportsmen 
and others objected to the bounty, and it was finally abolished in 
1893. By that time the bears were greatly decimated and in most 
provinces entirely exterminated. The building of railroads had con- 
tributed decidedly to this decrease. After considerable agitation 
for protection of the species, a new law of 1912 declared that no 
bear could be shot on crown land without special permission from 
the King, unless it had attacked man or domestic animals, nor on 
private land without the permission of the owner. According to a 
law of 1927, a bear killed anywhere belongs to the Crown. Com- 
pensation for damage by bears is now provided by the state. The 
whole sum thus paid for domestic animals (sheep, goats, reindeer, 
and one horse) , during 1933 and 1934 did not amount to more than 
2,404 kr. in all. Since the bears have been protected their numbers 
have increased only a little. 

In considering the bear's economic status, it may be noted that 
it never attacks man unless directly provoked or wounded. It preys 
on domestic animals only exceptionally, and not regularly. Many 
reindeer succumb to starvation and disease, and when the bear feeds 


on their carcasses, it is often accused of having killed the animals. 
Its diet consists largely of insects and their larvae and various kinds 
of plant food, especially berries. It also catches voles and lemmings. 
(Einar Lb'nnberg, in litt., October, 1936.) 

Under present conditions the bears are not threatened with ex- 
tinction (Einar Lonnberg, in litt., November 15, 1932) . 

Spain (U. a. pyrenaicus) . Cabrera (1914, p. 153) gives the range 
of this race as the Pyrenean and Cantabrian districts: Pyrenees 
of Aragon and Catalonia; mountains of Santander and Asturias; 
extreme north of the Provinces of Palencia and Leon and the 
eastern part of Lugo. In historic and even comparatively recent 
times it ranged more to the south, reaching at least the center of 
the peninsula. In 1582 Argote de Molina reported it not far from 

"Bears still occur not unfrequently all along the Cantabrian 
range of mountains. On the central chain of Spanish mountains 
they seem to be rarer. There are none now in Portugal. Formerly, 
as lately as the sixteenth century, before the devastation of the 
forests, the bear seems to have had a much "wider distribution in 
the Peninsula." (Gadow, 1897, p. 362.) 

In Asturias it nightly raids the maize-fields in the valleys in 
September. It is also in the habit of attacking and destroying many 
cattle. It is tracked to its covert, and a drive with beaters is organ- 
ized. From 20 to 30 bears are killed in Asturias every year. 
(Marquis de Villaviciosa de Asturias, in Chapman and Buck, 1910, 
pp. 296-297.) 

France (U. a. pyrenaicus in the Pyrenees; U. a. arctos in the 
Alps). Trouessart states (1884, pp. 195-196) that the species is 
restricted to the forested and the wildest regions of the Alps and 
the Pyrenees. It occasionally ravages the wheatfields and the vine- 
yards. It becomes more carnivorous with age and then forms the 
habit of making raids upon sheep and calves, and finally it even 
attacks grown cattle and horses. 

E. Bourdelle (1937, pp. 178-181, and in litt., March 6, 1937) gives 
the following account: 

Formerly rather widely spread in the mountainous regions of 
France Vosges, Jura, Cevennes, Alps, and Pyrenees it disappeared 
from the first three areas during the past century, and it now exists 
only in the Alps and the Pyrenees. It is generally believed to have 
disappeared from the French Alps and that the last two animals 
were killed in 1898 in the Forest of Vercors in these mountains. 
However, fresh tracks were observed in the same region in 1913, in 
1928, and again during the past few weeks. The extent and wildness 
of the Forest of Vercors militate in favor of the possibility of a 
few bears surviving there. 


In the Pyrenees the Brown Bear, though much less abundant 
than formerly, is still met with in the wild areas of upper Ariege 
adjacent to Andorra, in the Hautes Pyrenees (massif of Maladetta 
and Cirque de Gavarnie), and as far as the neighboring parts of 
the Basses Pyrenees (Forest of Irruti, for example). It is even 
probable that there are some bears in other parts. While it was 
still rather common 20 or 30 years ago, it has been gradually pushed 
back into refuges more and more restricted, where it is easily hunted, 
so that it has become rarer and rarer during recent years. 

Its only economic importance in the Pyrenees consists in its being 
a true game animal, its flesh being much prized in the whole region. 
Its hide provides a good fur, but not a very valuable one. 

The bear of the Pyrenees, like that of the Alps, is the victim 
of man's increasing penetration into the mountains, of the extension 
of agriculture, forestry, and mining, of highways, railroads, tourist 
traffic, and especially the sport associated with the hunting of this 

Protection should be provided for the Pyrenean bears as well as 
for those of the Alps (if the latter still exist in the Forest of Vercors) . 
Prohibition of hunting and of the sale of flesh and hides would 
suffice to halt the steady depletion and perhaps to assure the preser- 
vation of the species. 

According to Didier and Rode (1935, p. 268) , the last bears in the 
Hautes-Alpes were killed at the beginning of the nineteenth century. 
Two were killed in 1896 in the massif between La Chambre and 
Saint-Remy (Savoie) and Allevard (Isere). 

Belgium and The Netherlands. The species no longer exists in 
these countries (Martin, 1910, p. 38a; Ognev, 1931, p. 40). 

Germany. Blasius (1857, p. 199) reports the bear as still occur- 
ring in the Bavarian highlands. 

The last specimens were killed in 1759 in Thuringia, in 1770 in 
Upper Silesia, in 1810 in the Bavarian-Bohemian forests, and in 
1835 in Frauenstein, Bavaria. In the sixteenth century the species 
was still of frequent occurrence in Germany. When improved fire- 
arms were introduced, the population started a real war against the 
animal. The hide was readily sold, and bear-hunting, which in- 
volved some danger, was considered by the upper classes as enter- 
tainment and sport. Bounties were often paid, as the animals were 
looked upon in many places as seriously injurious to cattle. (Inter- 
nationale Gesellschaft zur Erhaltung des Wisents, in litt., October, 

The last bear in Pomerania was killed in 1750; in the Riesenge- 
birge about 1800. The species was still being taken in East Prussia 
up to 1806. (Krumbiegel, 1930, p. 6.) 


Castelli (1935, p. 33) quotes the Alpines Handbuch (1931) to the 
effect that the last one was killed in Wetterstein, Bavaria, in 1864. 

Denmark. The bear was once generally distributed over Den- 
mark. Remains have been found in the ancient kitchen middens, 
but there is no information on the occurrence of the species within 
historic times. (Winge, 1908, p. 127.) 

Switzerland. In 1869 Fatio (pp. 301-302) gave the following 
account of the bear's status: It was formerly abundant in the 
north and center of Switzerland but has gradually retired to the 
high Alps. It is now scarcely found except in Orisons, in Tessin 
(where 9 specimens were killed from 1852 to 1862), and here and 
there in the Jura. It has almost entirely disappeared in Valais 
and Uri. Basle, Lucerne, Schwyz, and Berne have no more bears. 

Castelli (1935, pp. 25-27) supplies the following records: The last 
bears near Zurich were killed or recorded in 1565, in Unterwald in 
1664, in Fribourg in 1698, while in Solothurn 38 were killed from 
1507 to 1737. Other last records are: Berne, 1815; Glarus, 1816; 
Vaud, 1843; Valais, 1860; Uri, 1898. In the Engadine 5 were killed 
in 1852, 8 in 1861, 6 in 1872, and 4 in 1873. In Orisons 25 were 
killed from 1878 to 1887, 9 from 1888 to 1897, and 3 within the 
following decade; the last one was killed in 1904 in Val Minger, but 
a female with two cubs was reported seen as late as 1919 in Val 

The species is now of exceptional occurrence in Switzerland, being 
represented only by an occasional straggler across the border from 
western Trentino, Italy (Tratz, in Castelli, 1935, p. 9). 

Italy. The bear is now restricted to two general areas in Italy 
the extreme north and the mountains of Abruzzi. 

From 1837 to 1852 146 specimens were killed in Trentino. The 
species is now protected there by the Italian Government. (G. 
Schlesinger, in Hit., March, 1937.) 

According to Castelli (1935, pp. 50-135), 77 bears were killed in 
Trentino from 1886 to 1912. In the district of Cles, at the north 
end of the Group of Brenta, the following numbers were killed: 26 
from 1886 to 1891; 5 in 1895; 2 in 1900; 1 in 1901; 2 in 1902; 4 in 
1903; 1 in 1906; 2 in 1908; 2 in 1909; 3 in 1910; 8 in 1911; 2 in 
1912; 3 in 1913. In Trentino 15 bears have been killed from 1922 
to 1933, and a small number have been seen yearly up to 1935. The 
Brenta Group and vicinity form the last refuge of the species in 
Trentino. It is sedentary there, and is in urgent need of protection, 
such as would be afforded by the establishment of a National Park. 

Castelli (1935, p. 28) quotes Cermenati to the effect that 40 bears 
were killed between 1876 and 1886 in Valtellina, Lombardy, Italy. 
He adds (p. 31) that the last individuals were killed in Valtellina in 
1896 and 1902. He also mentions (p. 32) a report of Depoli in 1928 


that there were about 10 bears, protected by law, in the Province of 
Carnaro, northeastern Italy. 

In the National Park of the Abruzzi about 200 Brown Bears are 
well protected (Tratz, in Castelli, 1935, p. 9). They are found in 
the mountains about the valley of the Sangro, and must be regarded 
as indigenous, notwithstanding the local tradition that the Czar of 
Russia had sent King Ferdinand of Naples a couple of such animals, 
which he set free in the mountains of the Abruzzi (Colosi, 1933, 
pp. 48-49). The park administration estimates the present number 
at about 100 (Laboratorio di Zoologia Applicata a Caccia, in Hit., 
September, 1936) . 

The bear is completely gone from the Sila Montains, Calabria, 
though present there in the middle of the last century (Hecht, 1932, 
p. 23). 

Austria. This was probably an indigenous species all over Aus- 
tria in former days. In Carinthia it was generally distributed up to 
1850; one bear was killed during each of the years 1895, 1920, 1927, 
and 1936. They are supposed to have come from the reserves in 
Gottschee, Carniola, and on Schneeberg (Monte Nevoso), north of 
Fiume, Italy ; perhaps also from Croatia. In Lower Austria the bear 
was observed rather frequently up to the last half of the nineteenth 
century ; here, in Semmering, Schneeberg, Rax, and the mountainous 
areas to the westward, fine stocks of bears were to be found. The 
last one was observed in 1919 near Rohr in the mountains of Lillien- 
feld. In Upper Austria and Salzburg the species was probably quite 
common up to the middle of the nineteenth century. In Tyrol the 
decrease started in 1570. At that time Duke Albrecht prohibited 
the capture and killing of bears. During the Thirty Years' War the 
numbers increased again. Up to about 1840 the annual kill was 
from 20 to 30 specimens. The last one was shot in Stellental, Tyrol, 
in 1898; in Vorarlberg, in 1870. The bear is not compatible with 
cattle-raising or with the increase in human population. (G. Schles- 
inger, in Hit., March, 1937.) 

According to the Alpines Handbuch (1931), 34 bears were killed 
in Tyrol in 1835, and in the same year the last one was killed in the 
Schneeberg district near Vienna. The last one was seen at Kar- 
wendel, on the Tyrolean-Bavarian border north of Innsbruck, in 
1896. (Castelli, 1935, p. 33.) 

Czechoslovakia. The species is still comparatively common in 
two well-defined districts. One embraces the mountainous territory 
of the Low and the High Tatra, bordered on the west by the Arva 
and the Waag Rivers, on the east by the Dunajec and Poprad 
Rivers. The other comprises the wooded Carpathians west of the 
railway from Munkac to Volovec. According to Dr. Komarec of 



Prague, 210 Brown Bears live in this territory, under government 
protection. (Tratz, in Castelli, 1935, pp. 8-9.) 

Hungary. Blasius (1857, p. 199) reports the bear as still occur- 
ring in the Hungary of his time, especially in the Carpathians and 
in the Hungarian Erzgebirge. The species is not found in the 
reduced Hungary (since World War I) (J. Schenk, in litt., Novem- 
ber, 1936). 

Yugoslavia. In this country the bear lives especially in the 
Gottschee district, Carniola, where it is carefully protected, and in 
the immense woods of the Auersperg district, Carniola (Tratz, in 
Castelli, 1935, p. 9). 

In Croatia about 20 bears are estimated to inhabit the forested 
area about Jasenak in the Grosse Kapela. They are also reported 
as not rare near Otocac and in the northern Velebit Mountains. The 
bears do far less damage than the wolves to livestock, and are 
reported as harmless to man. (Wettstein, 1928, p. 33.) 

The species occurs in considerable numbers only in Bosnia. It is 
found also in Slovenia (forests of Kocevje), in Croatia (forests of 
Velebit and Vemika Kapela mountains) , and in some parts of Serbia. 
In these regions a total of 272 specimens were killed from 1891 to 
1921, including 21 in 1892, 22 in 1893, and 26 in 1910. From 1921 
to 1931, 51 specimens were killed. (M. Hirtz, in litt., December, 

Albania. According to Baldacci (1932-33), the Bear still occurs 
commonly in the mountains in the center and north of Albania 
(Castelli, 1935, p. 37). 

Greece. The bear occurs in Macedonia and Epirus and does not 
show a decrease (Game Department, Ministry of Agriculture, Greece, 
in litt., October, 1936) . It is not a rarity in the extensive forests of 
the Greek and Turkish Balkans (Tratz, in Castelli, 1935, p. 9) . 

Bulgaria. "The bears are being killed at all seasons of the year, 
in every manner." A 220-kilo specimen was recently killed in a 
predatory animal "drive" in the Rhodope Mountains. Reserves 
for the preservation of the bear are advocated in the Eastern 
Balkans (Stara Planina) and in the Rhodope and the Pirene Moun- 
tains. (H. W. Shoemaker, in litt., June 30, 1932, and December 
27, 1932.) 

The species is found in all the mountains of Bulgaria. The present 
number is estimated at about 500. Since 1935 the bear may be 
hunted only on a special license, which is issued only for individuals 
that have become harmful to cattle pasturing in summer on the 
mountains. Protection is assured, and there are some reserves where 
hunting is entirely forbidden. (Bulgarian Game Association "Sokol," 
in litt., February, 1937.) 


Rumania. The bear occurs in the coniferous forest zone of the 
Banat and the southern Carpathians and in a limited area of the 
eastern Carpathians. Toward autumn it comes in search of food to 
lower heights, as far as the lower border of the coniferous forests 
(Tismana, in the Horjin District, and Brasov). (Calinescu, 1930, 
p. 365.) 

In the Transylvanian Alps it is still common in some districts 
(Tratz, in Castelli, 1935, p. 9) . 

Poland. The species was not rare in the eighteenth century, when 
it was still found all over Poland. The number is now reduced to 
about 250-270 individuals, found chiefly in the eastern Carpathians ; 
there are still about 20 in eastern Poland. There are also some in 
the Tatra Mountains. The number has perhaps increased of late. 
Females and young are absolutely protected, and the hunting of 
males is forbidden from January 15 to December 15. (M. Siedlecki, 
in Hit., October, 1936.) 

About 256 are left in the Carpathians, and 15 in eastern Poland 
in the swampy forest of Agarkow (National Council for Nature 
Protection, in litt., October, 1936) . 

In the future special permits for the shooting of bears will be 
granted by the Ministry of Agriculture (Quarterly Information 
Bulletin concerning the Protection of Nature in Poland, Kwartal 3, 

The proposed National Parks of the Tatra and of Czarnohora 
will be of importance in the protection of bears. The chief aim of 
the proposed International Park of Poland, Czechoslovakia, and 
Rumania is the creation of a breeding ground for the Bear, Lynx, 
Wildcat, Wolf, Stag, etc. (Benedyct Fulinski, MS, 1933.) 

Lithuania. The species has been exterminated in this country 
since about 1877 (T. Ivanauskas, in litt., November, 1936) . 

Latvia. The species is now extinct in Latvia, the last specimens 
having been killed in 1880-90 (N. von Transehe, in litt., February, 
1937). An occasional straggler comes from Russia or Estonia to 
our northeastern forests (Forest Department, Latvia, in litt., March, 

Estonia. About 20 individuals are found in the northeastern 
part, in the district of Wirumaa (Wiesland). Hunting is allowed 
only on a special permit from the State Forest Department. In 
recent years permits have been given for only one specimen each 
year. (Zoological Institute, University of Tartu, in litt., October, 

Russia. In Russia and Siberia the Brown Bear is more or less 
generally distributed, and in many areas it has maintained itself 
in fairly satisfactory numbers. Under these circumstances it seems 
unnecessary here to devote a great deal of space to the local dis- 


tribution. Very detailed information on this point is supplied by 
Ognev (1931, pp. 34-108). Various forms have been described from 
this vast region; all that are considered valid at all are rated as 
subspecies of Ursus arctos by Pocock (1932). 

W. G. Heptner writes (in litt., December, 1936) that the species 
is found in all forested regions of the U. S. S. R., including Caucasia 
and the mountains of Turkestan. In certain regions there are great 
numbers. Hunting is allowed in most regions during the whole year, 
but in White Russia only on special permit. In one part of Caucasia 
and in the mountains of Turkestan hunting is limited to certain 
open seasons. 

In European Russia, at the present time, the species seems to be 
found chiefly in the northern parts, in the Ural region, and in 
Caucasia. Many of the records from central Russia seem to date 
from the last century, and yet the species still survives near Lenin- 
grad and Moscow. In the Caucasus region generally it is quite 
common, though rare in Daghestan. (Ognev, 1931, pp. 34-38.) As 
many as half a dozen different races have been recognized in Cau- 
casia by various authors (Satunin, Smirnov, Lonnberg, Ognev) . 

"In the Caucasus, according to Prince Demidoff, it is so common 
that the keepers of the Grand Ducal territories have instructions 
to treat these animals as vermin, and to kill them whenever occasion 
occurs" (Lydekker, 1901, pp. 92-93). 

Asia. The Brown Bear is still numerous in many of the thickly 
forested areas of Siberia, where the people do not hunt so much now 
as formerly. It is distributed from the Urals east through the basins 
of the Ob, Yenisei, Lena, and Kolyma to the Anadyr region and 
Kamchatka (where it is very common). It ascends to 11,400 feet 
in the Sayan Mountains, and to 8,259 feet in the Yablonoi Moun- 
tains. (Ognev, 1931, pp. 38-40.) Southward its range extends to 
Turkey, Syria, Persia, Afghanistan, the Pamirs, Tian Shan, Hima- 
layas, western China, Manchuria, Hokkaido, and the Kuriles. There 
is almost a plague of bears in Hokkaido (Inukai, 19326, p. 526). 
Many different names (generic, specific, and subspecific) have been 
applied to the Brown Bears of various parts of Asia, but Pocock 
(1932) regards them as nothing more than races of Ursus arctos. 
Separate accounts of two of these forms follow. 

The Old World Brown Bear is closely related to the Grizzly Bears 
of North America and shows a decided resemblance to them in 
food habits and economic status. The considerable human tolerance 
exhibited toward it, together with its survival to the present day 
in most of the thickly populated countries of Europe, leads one to 
question the actual necessity for the ruthless war of extermination 
that has been waged upon the Grizzlies in the relatively sparsely 
settled areas of the Western United States. 



Manchurian Black Bear; Manchurian Grizzly 


Ursus lasiotus J. E. Gray, Ann. Mag. Nat. Hist., ser. 3, vol. 20, p. 301, 1867. 

("North China.") 

SYNONYM: Melanarctos cavijrons Heude. 
FIG.: Sowerby, 1923, pi. 3. 

For the purposes of the present report, this subspecies is restricted 
to Manchuria and adjacent regions of the Asiatic mainland, although 
Pocock (1932, p. 799) provisionally includes with it the bear of Yeso 
(Hokkaido) and the Kuriles (U. a. yesoensis Lydekker). 

Only five museum skins of this little-known bear seem to be on 
record from Manchuria and Mongolia (Pocock, 1932, p. 799). It is 
becoming increasingly rare, and calls for government protection in 
some way or other if it is to be saved from ultimate extinction 
(Arthur de C. Sowerby, in litt., April 24, 1937) . 

It is as large as the Kamchatkan Brown Bear (U. a. beringianus 
Middendorff) but differs from it on the average, at least, in the 
prevalent blackness of its hue. The general color is glossy black; 
muzzle brown ; underwool brown. Adult male from Manchuria : head 
and body, 6 feet 7 inches; tail, 5.5 inches (Pocock, 1932, pp. 799- 

The range seems to include the forested regions of northern 
Manchuria, northern Mongolia, southeastern Siberia, and perhaps 
northern Korea. 

Sowerby (1920, pp. 230-231) shot a specimen in North Kirin, 
Manchuria, and heard reports of a similar animal in South Kirin, 
on the lower Sungari River, and in northern Korea. "The specimen 
I shot was very savage .... The native Russians and Chinese 
greatly fear this animal, as it has been known to kill and devour 

Sowerby also writes (1923, p. 58) : "The distribution of this 
species is doubtful, or, perhaps it would be more correct to say, is 
not known. So far it has been recorded only from the forest near 
Tsi-tsi-har in South-western Heilung-kiang, and from the forest 
in the I-mien-p'o district of North Kirin. From all accounts, how- 
ever, it occurs throughout the Manchurian forest, and on into 
Primorsk [Siberia]." He adds that a hunter reports this form as 
"much rarer than the black bear [Selenarctos] , occurring in the 
proportion of one in twenty of the bears shot in the district." 

Syrian Bear 

URSUS ARCTOS SYRIACUS Hemprich and Ehrenberg 

Ursus syriacus Hemprich and Ehrenberg, Symbolae Physicae Mammalium, 
decas prima, text to pi. 1, 1828. (Near the village of Bischerre, Mount 
Makmel, Lebanon.) 


SYNONYM: Ursus schmitzi Matschie (1917). 

FIGS.: Hemprich and Ehrenberg, 1828, pi. 1; Wolf, 1861, pi. 17 (specimen 
from the Persian Gulf) ; Pocock, 1937, p. 807, fig. 

The Syrian Bear is now extinct, or nearly so, in Palestine and 
Lebanon but survives in rather indefinite numbers to the northward 
and eastward, where the exact limits of its range have not been 

The type specimen (which was not full-grown) was described as 
uniform fulvous-white; it was smaller than Ursus arctos and had 
long ears. Its head and body measured 3 feet 8 inches; tail, 6 
inches ; height at shoulder, 2 feet 4 inches. Other skins were said to 
be fulvous or sometimes almost wholly brown. (Hemprich and 
Ehrenberg, 1828.) 

The range, according to Flower (1929, p. 149), is "western Asia: 
in certain mountainous localities from Asia Minor and Syria to 

Bodenheimer (1935, p. 114) writes: 

The Syrian Bear . . . was not uncommon in N. Palestine in Biblical times. 
David boasts of having strangled a bear, which had attacked his herd (I Regum 
17, 34) and two bears killed the 42 boys who had scoffed at the prophet 
Elisha (II Regum 2, 24). Tristram encountered one in a ravine near Tiberias, 
near Beisan and in the Jolan. Schmitz seems to have seen the last specimens on 
the southern Hermon (1911, 1913). ... It has not been a menace to flocks of 
sheep and goats for a long time, but occasional visits to vine-yards and 
fruit-groves are still reported from Syria. The Bear is extinct on the Hermon 
and Anti-lebanon, mainly because it was so drastically hunted by German 
officers during the war. It is reported to have survived on the Lebanon. 

J. C. Phillips writes (in litt., July 20, 1936) that there were 
supposed to be a few bears left on Mount Hermon when he was 
there in 1912. 

The following information, supplied by Dr. William Van Dyck 
and Professor West, both of the American University in Beirut, is 
transmitted by Theodore Marriner (in litt., 1936) : 

"Shortly after the World War, when there were a large number of 
army rifles in mountain villages, the number of Syrian bears . . . 
was greatly reduced. They were, in fact, exterminated in some parts 
of the Anti-Lebanon range, but a few are still reported in the less 
accessible parts of both the Anti-Lebanon and Lebanon ranges. 
Farther north, in the Gebel Ansariyah and in the Amanus range of 
northern Syria and southern Turkey, they are still quite common 
in the more wooded sections. At the present time no definite attempt 
is being made to preserve the Syrian Bear, although the government 
policy of forbidding civilians to carry rifles indirectly helps towards 
this end." 

Aharoni (1930, pp. 336-337) gives the following account (some- 
what freely translated) : "During the war, while stationed in Leba- 


non, I found that the light Isabella-colored bear [Ursus syriacus 
H. and E.] , with the dimensions of the original description, inhabited 
only the green shrubbery of the Anti-Lebanon, while the smaller 
brown bear [U. syriacus schmitzi Matschie] inhabited only the bare 
snow-fields of the Lebanon. I saw examples of both subspecies in 
nature and still have specimens from Lebanon. To-day the bear 
has disappeared not only from Palestine, but perhaps also from 

"Last year I became convinced that the Mesopotamian bear in- 
habiting the Jebel Abdul-Aziz [in the present Syria; lat. 36 30' 
N., long. 40 30' E.] represents a distinct subspecies." 

F. S. Bodenheimer writes (in Hit., March, 1937) that the animal 
is now extinct in Palestine and Lebanon but probably still survives 
in Anti-Lebanon in small numbers. He adds that protection is most 
highly desirable. 

Pocock states (1932, p. 793) that "the bears of Asia Minor and 
Syria merely differ from the typical Brown Bear of Europe in being 
on the average paler in colour, intermediate specimens occurring 
in the Caucasus and perhaps in northern Persia." He records speci- 
mens from Smyrna and from Sumela, 30 miles south of Trebizond, 

Blanford (1876, pp. 46-47) gives the following account of bears 
in Persia: 

"Major St. John, . . . who has seen several Elburz bears, assures 
me that, although they are darker than the true Ursus Syriacus 
which is found in Southern Persia, they are much paler in colour 
than the common bear of Europe. . . . ' 

Ursus syriacus "is, as Major St. John assures me, the bear of 
South-western Persia. It is not the bear of Baluchistan, but is said 
to be found between Bampur and Bam. It is found* pretty commonly 
in the neighbourhood of Shiraz and in the hills bordering on Meso- 

To this St. John adds (in Blanford, 1876, p. 47) : "This bear is 
found throughout the mountains of Western and Northern Persia, 
possibly extending to Khorassan. In many places watchers are set 
at night to keep the bears from the ripening grapes." 

Atlas Bear; Crowther's Bear 


Urs[us~\ Crowtheri Schinz, Synopsis Mammalium, vol. 1, p. 302, 1844. (Based 
upon "the Bear of Mount Atlas," Blyth, Proc. Zool. Soc. London 1841, 
p. 65; type locality, "the foot of the Tetuan mountains, about twenty-five 
miles from that of the Atlas.") 

The bear of North Africa is almost a mythical species, for no 
specimen has ever reached a museum. No very definite news of the 


species has been obtained for nearly a century, and it is very prob- 
ably extinct. 

An adult female was smaller than the American Black Bear but 
more robustly formed and with a shorter and broader face, though 
the muzzle was pointed; toes and claws remarkably short; hair 
black or brownish black, shaggy, 4 or 5 inches long; muzzle black; 
under parts orange-rufous (Blyth, 1841, p. 65) . 

Pliny, though skeptical himself, quoted Roman annals to the 
effect that Domitius Ahenobarbus, an aedile of 61 B. C., had shown 
in the Roman arena a hundred Numidian bears, conducted by as 
many Ethiopian hunters. The bear of Libya was mentioned by 
Pliny's contemporaries, Juvenal and Martial, and a long time pre- 
viously by Virgil. (G. Cuvier, 1825, vol. 4, pp. 325-326.) 

Strabo says expressly that the Moors dressed themselves in bear 
and lion skins (Wagner, 1841, p. 70). 

Blyth (1841, p. 65) gives a brief description of the animal, based 
upon information supplied by Mr. Crowther, who had spent some 
time in Morocco. "Upon questioning Mr. Crowther respecting the 
Bear of Mount Atlas, which has been suspected to be the Syriacus, 
he knew it well, and it proves to be a very different animal. . . . 
This individual was killed at the foot of the Tetuan mountains, 
about twenty-five miles from that of the Atlas. It is considered a 
rare species in that part, and feeds on roots, acorns, and fruits. Does 
not climb with facility; and is stated to be very different-looking 
from any other Bear." An unsuccessful effort was made to preserve 
the skin of the specimen mentioned. 

According to Loche (1867, p. 52), Shaw (1743) mentions a bear 
in the Atlas Mountains. Loche also states that the Emperor of 
Morocco had recently sent to the zoological garden of Marseilles a 
live bear coming from his territory. 

Bourguignat (1867, pp. 41-46) contributes the following informa- 
tion. Herodotus records a bear from western Libya. Poiret, a French 
botanist and zoologist, reports (1789) bears from the Atlas Moun- 
tains, and mentions a fresh skin brought by an Arab into Mazoule. 
A friend of Bourguignat's, M. Letourneux, had reports of many 
bears in the region of fidough, and learned of others occurring not 
long previously on Djebel-Bou-Abed, Djebel Gherar, Djebel Debhar, 
and Djebel Thaya, Algeria. The animal was said to be small, thick- 
set, and brown, with a white spot on the throat, and to be very fond 
of honey and fruits. Bourguignat himself records skeletal remains 
of a bear from a cavern on Djebel Thaya in the Province of Con- 
stantine, to which he gives the name of Ursus jaidherbianus. Human 
artifacts associated with these remains were believed to date from 
the early Christian Era. 


Lataste (1885, pp. 235-237), in reviewing the evidence for the 
presence of bears in Barbary, considers that the case has by no 
means been proved. 

"Since Mr. Crowther's time no more definite news has been re- 
ceived of this bear, though other travellers have reported statements 
of Arabs and Moors that such a creature exists in the mountains 
of Eastern Morocco and Western Algeria" (Johnston, in Bryden, 
1899, p. 608). 

"In view of the apparent rarity of the animal, it is important 
to mention that fossilised remains of bears have been discovered in 
caverns in north-western Africa, as well as in the rock-fissures of 
Gibraltar" (Lydekker, 1908, p. 463). 

During the years 1892-96, "fchere were still rumours of Bears 
(Ursus crowtheri) in the Western Atlas, but although they certainly 
existed there in the first half of the last century I have never heard 
of one being killed or seen since this region became better known 
with the penetration of the French into Morocco, though there may 
be a possibility that a few exist" (Pease, 1937, p. 81). 

The foregoing accounts seem to constitute fairly strong evidence 
of the former existence of a bear in North Africa. It must be 
acknowledged, however, that no less an authority than Cabrera 
(1932, pp. 10, 102-103) throws the whole case for the Atlas Bear 
out of court. But he is hardly correct in maintaining that its sole 
basis is the "fantastic" account of Blyth. 

On first thought, the Atlas Bear might appear to be a note- 
worthy exception to the general rule that recently extinct mammals 
have succumbed to the advance of the European type of civilization. 
Yet one of the tools of that civilization, the rifle, in the hands of 
the Moors, must have at least contributed to the animal's downfall. 
Nevertheless, the disappearance of the Barbary Lion from Morocco 
in the early part of the present century is singular enough (Cabrera, 
1932, p. 186) , and the still earlier disappearance of the Atlas Bear 
is even more puzzling. 

Family MUSTELIDAE: Weasels, etc. 

This family is distinguished by the large number of valuable fur- 
bearers represented in it; and many of the species have been seri- 
ously reduced by the demands of the fur trade. Its distribution is 
practically as cosmopolitan as that of the Canidae; it extends to 
Borneo and the Philippines, but not to Australia. There are about 
35 genera and 400 species and subspecies. Nineteen forms (including 
one extinct species) are discussed in Dr. Allen's volume on the New 
World (1942), and nine forms of the Old World in this volume. 


European Mink. Vison (Fr.). Norz; Sumpf otter (Ger.) 


[Mustela] Lutreola Linnaeus, Syst. Nat., ed. 12, vol. 1, p. 66, 1766. (Finland.) 
FIGS.: Royal Nat. Hist., vol. 2, p. 68, 1894; Martin, 1910, pi. 33; Didier and 
Rode, 1935, p. 303, fig. 176. 

The European Mink is a rare and vanishing species in central 
Europe and France but is more generally and more commonly dis- 
tributed in Russia. 

The general color is a rich dark brown; region about the mouth 
whitish; tip of the tail blackish. Head and body, 350 (female) to 
400 mm. (male); tail, 130 (female) to 140 mm. (male). (Miller, 
1912, pp. 415, 418.) 

The range extends from western France eastward to the Tobol 
and Irtish Rivers in western Siberia; south to Austria, Hungary, 
Rumania, and Transcaucasia ; north to Finland and northern Russia. 

France. Though recorded by Lesson in 1840 in Poitou and Sain- 
tonge, the Mink was long overlooked in France. It seems to have 
been formerly rather common in the center, the west, the southwest, 
Normandy, and the Vosges. The present range consists of a narrow 
zone extending from the Jura to the vicinity of Nantes and in a 
general way following the valley of the Loire. Here the species 
seems to become rarer and rarer. The decrease is due to the active 
hunting of the animal, for its fur is very valuable and it is also 
considered a harmful species. Prohibition of hunting and surveil- 
lance of the fur trade would be the only means of conservation; 
but these measures would be very difficult to apply. (E. Bourdelle, 
in litt., March 6, 1937.) 

Martin (1910, p. 35a) extended the range to the Gironde and to 

The Mink's food includes fish, frogs, crawfish, ducks, and small 
mammals (Didier and Rode, 1935, p. 304) . 

Germany. The species has been exterminated in western Ger- 
many and is very rare in northern and eastern Germany. Latest 
dates of capture are: Mecklenburg, 1894-96; Hannover, 1902; East 
Prussia, 1909; Liineburger Heide, 1910. At present there is no open 
season. (Internationale Gesellschaft zur Erhaltung des Wisents, in 
litt., October, 1936.) 

It had disappeared from Schleswig-Holstein by about 1890 (Mohr, 
1931, p. 32). 

During recent years solitary individuals are still regularly shot or 
seen in the east (Krumbiegel, 1930, p. 6) . 

Switzerland. Fatio (1869, p. 336) has only a few doubtful records 
from this country. 

Austria. The Mink was formerly found in Burgenland and prob- 



ably also in Lower Austria. It is now absolutely protected in Bur- 
genland, where it is said to still exist, though reliable reports are 
not obtainable. (G. Schlesinger, in litt., March, 1937.) 

Czechoslovakia. The known specimens are mostly from the 
Carpathians, in the former Hungary (J. Schenk, in litt., November, 

Hungary. From the present limits of Hungary (since the World 
War) only one specimen is known. The animal has no legal pro- 
tection. (J. Schenk, in litt., November, 1936.) 

FIG. 23. European Mink (Mustela lutreola). After Lydekker. 

Rumania. The Mink is common, like the Otter, but has a greater 
distributional area, since it is more adaptable to civilization (Cali- 
nescu, 1930, p. 366) . 

Poland. It was formerly quite common all over Poland but is 
now very rare, being found especially in the southeast, in Polish 
Podolia and in the Eastern Carpathians. Hunting is forbidden 
from February 1 to December 31. Lately there has been a demand 
for complete protection. (M. Siedlecki, in litt., October, 1936.) 

Kuntze (1935, p. 63) records it from northeastern and south- 
eastern Poland. 

Lithuania. The species is exterminated except in the eastern 
part of the country, where it is still found in the districts of Zarasai 
and Utona. The annual production amounts to as many as 150 
skins. The value of one is about 40 Litas (5-6/) . So far no protec- 
tive measures have been adopted. (T. Ivanauskas, in litt., Novem- 
ber, 1936.) 

Latvia. In 1908-09 it was reported as numerous in Courland 
and widely distributed in Livonia (Ognev, 1931, p. 759). 


The present stock is about 2,000 individuals. The increase through 
natural propagation is not important, and the annual kill is about 
300-400 specimens. The species is protected from March 1 to Novem- 
ber 14. Forest guards are not allowed to kill it. (Forest Department, 
Latvia, in litt., March, 1937.) 

Estonia. The species is found throughout the country in suitable 
areas. There is a steady decrease, owing to the drying up of the 
country. The animal is not threatened by man, but it is without 
any legal protection. (Zoological Institute, University of Tartu, 
in litt., October, 1936.) 

Finland. The species is apparently distributed in the southern 
part of the country (Ognev, 1931, p. 758). 

Russia. From Ognev's data (1931, pp. 758-761), the Mink ap- 
pears to be widely and more or less commonly distributed over most 
of Russia, from Kandalaksha Bay, the lower Dvina, and the Pet- 
chora and Usa Rivers in the north to the Ukraine, the Caucasus, 
and Astrakhan in the south. Westward it is found about Lake Onega, 
in Volhynia and Podolia, and on the lower Dniester. At the bazaar 
of White Russia 473 Mink skins were sold in 1926-27, and 649 in 
1927-28. Eastward the species is found in the Ural region, from 
the tributaries of the Petchora in the north to Orenburg and the 
Ilek River in the south. Beyond the Urals it extends only to the 
Tobol and Irtish Rivers. In Transcaucasia it occurs on the Bzyb 

The animal is strongly persecuted as a fur animal and is rare 
in certain regions. Hunting is not allowed in the Volga region and 
in the eastern part of European Russia. There is no danger of ex- 
tinction, except in certain industrial regions. (W. G. Heptner, in 
litt., December, 1936.) 

Russian Sable. Marte zibelline (Fr.). Zobel (Ger.) 


Mustela zibellina Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 46, 1758. (Northern 

Asia; type locality restricted by Ognev (1925, p. 276) to "the northern 

part of the government of Tobolsk.") 
FIGS.: Royal Nat. Hist., vol. 2, p. 55, 1894; Ognev, 1931, pi. 5 (M. z. 

sahalinensis) ; Zeitschr. f. Saugetierk., vol. 9, pi. 18, fig. 7, 1934 (Amur 

form) . 

This is one of the animals that has suffered particularly from 
the "curse of beauty." It has been decimated by the demands of 
the fur trade and has disappeared from considerable areas within 
its former range. Its principal home is in Siberia. 

The Sable bears considerable resemblance to the Pine Marten 
(Martes martes). It has a cone-shaped head, large ears, a bushy 


tail, and comparatively stout limbs. The fur is thick and soft; the 
color varies from blackish, mixed with gray and brown, to yellowish 
brown; throat sometimes orange. Head and body, about 20 inches; 
tail, about 7 inches. 

The former range of the species included the forested regions 
from northern Russia east to the Anadyr district, Siberia; it ex- 
tended south to the southern Urals, the Altai and Sayan Mountains, 
Manchuria, the Ussuri district, Hokkaido, Sakhalin, and Kamchatka. 

The following subspecies have been recognized (cj. Ognev, 1925, 
and 1931, pp. 560-598) , but the ranges so far assigned to them do 
not cover the entire range of the species: 

Martes zibellina zibellina (Linnaeus). (Original reference and 
type locality given above.) Tobolsk Sable. 

Color dull and pale, varying from cinnamon-drab to pale brownish 
yellow and even to dark brown; underfur light and dull. 

Range: the Ob Basin and the Ural region. 

M. z. yeniseensis Ognev, Jour. Mammalogy, vol. 6, p. 277, pi. 26, 
fig. 3, 1925. ("Krasnoyarsk district, the forest on the plain along 
the Yenisei River," Siberia.) Yenisei Sable. 

Color more dusky warm brown in comparison with the Tobolsk 

Range: the great forests of the districts of Krasnojarsk, Ashinsk, 
and Kansk, in the Yenisei Basin. 

M. z. sajanensis Ognev, Jour. Mammalogy, vol. 6, p. 278, 1925. 
("Orsyba River, northern part of the Sajansky Mountains," Siberia.) 
Sayan Sable. 

General color dark brown; underfur pale yellowish. 

Range: "the mountain country of the rivers Uda, Kasyr and 
especially of the Kasyr-Suk and partly of the Usa." 

M . z. princeps (Birula) . (Mustela zibellina princeps Birula, Ann. 
Mus. Zool. Acad. Imper. Sci. Petrograd, vol. 22, p. 08, 1922; the 
mountain country of Bargusin, Transbaikalia, Siberia.) Bargusin 

Fur soft and silky; color a brilliant blackish brown; underfur 
bluish gray, brownish at bases and tips; throat patch much reduced, 
commonly not visible. 

Range: mountain forests, Bargusin Hills and spurs of the Stano- 
voi Mountains, Transbaikalia. 

M. z. kamtschadalica (Birula). (Mustela zibellina subsp. kamt- 
schadalica Birula, C. R. Mus. Zool. Acad. Sci. Petrograd 1918, 
p. 82 (fide Ognev) ; Kamchatka.) Kamchatka Sable. 

General color between warm sepia and mars brown; underfur 
pale yellowish gray ; skull large. 

Range: Kamchatka. 

M. z. sahalinensis Ognev, Jour. Mammalogy, vol. 6, p. 279, pi. 26, 
fig. 4, 1925. ("Saghalien, Wedernikovo.") Sakhalin Sable. 


Winter pelage like that of the Kamchatka Sable but of a more 
decided cinnamon tint and lighter; throat patch of the same cinna- 
mon color; head avellaneous, back darker; flanks sayal brown or 
tawny-olive; underfur pale yellowish, more cinnamon at the tips; 
summer pelage duller and darker, more brownish. 

Range: the whole of Sakhalin Island. 

M. z. brachyura (Temminck). (Mustela brachyura Temminck, 
in Siebold, Fauna Japonica, Mammiferes, p. 33, 1844; Matimaja, 
Hokkaido, Japan.) Japanese Sable. 

FIG. 24. Russian Sable (Maries zibellina subsp.) 

Inferior to the Siberian Sable in fineness and length of fur; back 
and tail dark brown; sides and limbs lighter; long hair of feet con- 
cealing the claws. Tail, 3.5 inches. (Temminck, 1844, pp. 33-34.) 

Range : Hokkaido and the Kuriles. 

Russia. In past centuries the Sable's range extended westward 
perhaps as far as the Kola Peninsula or even Lapland. In the six- 
teenth and seventeenth centuries it was found on the Petchora 
River and probably at the same time in the Dvina region. In Pallas's 
time it occurred in the vicinity of Ufa, west of the southern Urals; 
the last one in this general region was killed in 1850 near Ufimsk. 
Its southern limit on both slopes of the Urals was about latitude 
52 N., or possibly 51 N. About 1700 it inhabited the entire Gov- 
ernment of Perm and the eastern half of the Governments of Vo- 
logda, Archangel, and possibly Viatka. By 1875 about 300 Sables 
were trapped annually in the northeastern part of Perm. More 


recently the Russian population of the species seems to have become 
largely restricted to the Urals, where it is less numerous on the 
western slopes than on the eastern. However, in 1925 it was still 
common on the Ilych River, a tributary of the Fetch ora. (Ognev, 
1931, pp. 569-570.) 

The Ural form is considered distinct from M. z. zibellina but is 
not named. It occurs sporadically and rarely along the western 
foothills (headwaters of the Shugora, Laga, Ilych, and Petchora) ; 
it is also rare on the eastern slopes, at the sources of the Losva, 
Aspia, Purma, Ushma, Toshemok, and Wishaj Rivers. The fur is 
considered the finest in western Siberia. (Ognev, 1925, p. 277.) 

Siberia, Ob Basin (M. z. zibellina). In the Government of To- 
bolsk the Sable is not rare in the taiga forests of the Pelym River; 
it is rare along the Tavda River and in the Tarsk and Surgut dis- 
tricts. At the end of the last century more than 300 Sables were 
obtained annually along the Jugan River. The species is absent 
between Beresof and Obdorsk. Along the Rivers Omi, Tara, and 
Irtish, and in the southern part of the Government of Tobolsk it 
was very scarce in 1886. In the Narym district it was numerous on 
the upper Wasugan River in 1875. It occurs on the Ket and Chulym 
Rivers, and was particularly numerous on the Tchirk-Ul River about 
1923. It avoids the steppes in the central part of the Government of 
Tomsk. (Ognev, 1925, p. 277, and 1931, pp. 571-572.) 

Prejevalsky (1879, p. 233) reported the Upper Katuna, the Bukh- 
tarma, and their tributaries, in the Russian Altai, as particularly 
good districts for Sables. The hunters used specially trained dogs, 
and endeavored to surround the animals with nets, which were as 
much as 1,000 feet long and 4 feet high. The average price of a 
sable skin was then 15 rubles. 

Siberia, Yenisei Basin (M. z. yeniseensis, M. z. sajanensis, M. z. 
princeps) . The Sable is found in suitable areas from the Mongolian 
boundary northward to latitude 69 N. The form living along the 
Tunguska River and near Turukhansk probably represents an un- 
described subspecies ; the same form is found in small numbers in the 
adjacent Khatanga Basin. The species is less common in the Gov- 
ernment of Irkutsk than in the Government of Yenisei. It is absent 
from the steppes in the vicinity of Minusinsk, Achinsk, and Kras- 
noyarsk. It is common in the Sayan region on the Kasyr-Suk and 
Uda Rivers, and occurs on practically all sides of Lake Baikal. 
According to Turov (1923) , 700 skins were exported annually across 
Bargusin from the Verkhne Angarsk and Podlemorsk districts near 
Lake Baikal. (Ognev, 1931, pp. 572-573.) 

Siberia, Lena Basin. In the Olekma-Vitim mountainous country 
the Sable is very rare. Far to the north, in the enormous region 
between the Anabar, the Olenek, the Lower Tunguska, and the Vilui, 


it disappeared about the middle of the last century. About 1873 
it was found occasionally on the Patom River. Its disappearance 
from the entire southern Muisk district was unusually rapid. It 
has been recorded from the Aldan and Mae Rivers. (Ognev, 1931, 
p. 573.) 

The dark animals furnish the most valuable skins; in peace times 
they fetched as much as 2000 marks. They come mostly from the 
Vitim Plateau and from the Bargusin district, where the Sable is 
now almost exterminated. (Klemm, 1930, p. 367.) 

Eastern Siberia. At present the Sable is not found in the Verk- 
hoyansk and Kolyma districts. In former times it was widely 
distributed along the Kolyma and Omolon Rivers, but it finally 
disappeared from the Kolyma district in 1852. The species has 
long since vanished from the Anadyr River region ; the last one was 
found near the village of Eropol about 1847. 

By 1900 the species was rare in the Gizhiginsk district, though 
in former years from 30 to 50 Sables were collected annually, par- 
ticularly from the Penzhina Valley and from northern Kamchatka. 
Possibly this form belongs to M. z. kamtschadalica, which is widely 
distributed in Kamchatka, especially in the Petropavlovsk district. 
(Ognev, 1931, pp. 574-575, 595.) 

In Kamchatka the Sable was decimated in Dybowski's time (1879- 
85). At the beginning of the nineteenth century a hunter could 
get 40 animals a day, and the annual production of Kamchatka 
amounted to 10,000 skins. The natives did not endanger the stand 
of Sables, but by 1881 Cossack and Tungus immigrants reduced the 
yield to 2,883. The abundance of the animals in some years was 
dependent on the wholesale occurrence of a vole, Microtus oecono- 
micus. (Kuntze, 1932, p. 47.) 

In the western Amur region the species occurs on the Argun and 
Shilka Rivers. In the middle and lower Amur Basin, the Sables 
from the Albazin area, the Zeya River, and the Bureya Mountains, 
which are very dark in color, and costly, may belong to the sub- 
species M. z. princeps. In 1861 the species was reported as particu- 
larly numerous on the Amgun River. In the Ussuri district it varies 
from common to rare, and has even disappeared entirely in some 
parts. The Ussuri Sable is probably very near to M . z. sahalinensis. 
(Ognev, 1925, pp. 279-280, and 1931, pp. 573-574.) 

Sowerby (1923, pp. 63-65) says that among the Tartars of the 
Primorsk coast in southeastern Siberia, "sable hunting is their chief 
end and aim in existence." He continues: 

It is certain that it was largely the presence of the sable throughout Siberia 
and in the Amur and Primorsk that led the Russian pioneers and conquerors 
across that wide stretch of country. . . . Thus we must look upon this little 
animal as having a very important bearing upon the history of these regions. 


. . . The only trouble is that with the unrestricted hunting that takes place, and 
the steady increase in the settlement of the country, this valuable supply of 
fur-bearing animals is rapidly diminishing. . . . 

It is evident that one of the chief objects of the conquest of Siberia was 
to secure a supply of sable skins for the Imperial Government, and it is 
significant that the conquering Cossacks . . . always imposed a heavy tribute 
of sable skins upon the Tartar tribes they defeated, and brought under 
subjection .... 

In Siberia this animal is protected by the Government, and comparatively 
recently it was given a five years closed season. 

The Cedar Valley Reservation (Kedrovaya Pad) on Amur Bay, 
comprising 7,500 hectares, and the Kronotsk Bay Reservation in 
Kamchatka, comprising 15,000 hectares, provide for the protection 
of the Sable (Makaroff, in Skottsberg, 1934, pp. 433-434) . 

In Sakhalin the Sable is distributed over the entire island, and in 
1889 it was considered more numerous there than in any other part 
of Siberia (Ognev, 1931, p. 574). 

Mongolia. P. P. Sushkin reported in 1925 that the Sable was a 
regular inhabitant of the southern slopes of the Altai Mountains, 
about the headwaters of the Black Irtish and the Urungu. It is also 
abundant in the vicinity of Kossogol, at the southern base of the 
Sayan Mountains. (Ognev, 1931, p. 572.) 

Manchuria. The most valuable fur-bearing animal of Manchuria 
is the Sable (Sowerby, 1934, p. 286) . 

"The Manchurian sable does not come up to those from the Amur, 
Primorskaya and Siberia in the value of its pelt. ... It is said 
that the Chinese nearly always hunt the sable by running it down 
with dogs. . . . 

"Unfortunately the Chinese Government is not alive to the value 
of its game and fur-bearing animals and birds, and so affords no 
manner of protection. There can be only one result of this; com- 
plete extinction of the sable in the provinces of Heilungkiang and 
Kirin." (Sowerby, 1923, pp. 64-65.) 

Japan. Temminck (1844, pp. 33-34) described the Japanese 
Sable from Yezo (Hokkaido) and added that it was common in all 
the Kuriles. It was being utilized in the fur trade in his day. 

In Hokkaido "the sables . . . have . . . met a sad fate and in 
spite of particular protection, their coming back to their existence 
as before is anything but promising. . . . 

"It is interesting to note that the increase of the Japanese minks 
[Mustela itatsi] in Hokkaido associates closely with the decrease 
of the sables which occupied the land before the minks and de- 
creased inland gradually from the southern part. The number of 
sables caught in Hokkaido was 2,395 in 1906, 765 in 1910, 1,706 in 
1915 and 214 in 1919 respectively." (Inukai, 19326, pp. 524, 527.) 

Uchida reports (1935, p. 8) a total of 5,948 Japanese Sables taken 
during the six-months open season of 1929-30. 


In the Kuriles the Sable is threatened with destruction (Miyoshi. 
in Skottsberg, 1934, p. 412) . 

Economics and conservation. The Sable has been very actively 
hunted for several centuries. Its numbers are now much reduced, 
and in only a few regions can it be said to be "not rare." Its range 
also has been considerably reduced and has become discontinuous. 
There has been shrinkage of the range at its western limits in Russia 
and at its southern limits in western and southern Siberia. From 
time to time hunting has been forbidden in the whole or in certain 
parts of its range. These measures have given good results, and the 
decrease in numbers has been halted for several years. In order to 
obtain an increase, a closed season has been maintained on the 
whole territory of the U. S. S. R. Several great reserves have been 
created to afford protection to the Sable (Barguzinsk, Kronotski, 
Sikhote-Alin, Kondo-Sosva) . Successful propagation has been car- 
ried out, and several "sovkhoz" have been specially created for 
sable-farming. (W. G. Heptner, in litt., 1937.) 

In 1928 the entire Sable production of Asia was 15,000. In 1929-30 
the yield in that part of eastern Siberia bordering Manchuria was 
1,925 skins (Kuntze, 1932, p. 47). 

Wolverine; Glutton. Glouton arctique (Fr.) Vielfrass (Ger.) 

GULO GULO (Linnaeus) 

[Mustela] gulo Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 45, 1758. (Lapland.) 
FIGS.: Fitzinger, Bild.-Atlas, Saugth., fig. 70, 1860; Royal Nat. Hist., vol. 2, 

p. 71, fig., 1894; Zeitschr. fur Saugetierk., vol. 8, pi. 31, 1933; Oguev, 

1935, pi. 2. 

The range as well as the numbers of the Wolverine have become 
reduced in northern Europe, but its status in northern Asia has 
remained more satisfactory. 

The general form is heavy and badgerlike; fur long and dense; 
tail bushy; general color a rich dark brown, becoming blackish on 
legs, feet, and tail ; a broad light brownish or yellowish band across 
rump and upper side of basal part of tail, extending forward to 
shoulders, where it gradually disappears; a cream-buff area across 
the head between ears and eyes. Head and body, 825 mm.; tail, 
125 mm. (pencil, 75). (Chiefly from Miller, 1912, pp. 434-440.) 

"In prehistoric times, the wolverine was found in England, and 
indeed ranged as far south as the Pyrenees" (Lydekker, 1901, 
p. 112). 

Its recent range is given by Miller (1912, p. 434) as the "northern 

forests of the Old World; in Europe, confined to Scandinavia and 

northern Russia." It is stated by Trouessart (1910, p. 71) a little 

more fully: "Circumpolar Europe (but not the islands north of the 



continent), south to lat. 55 N. (Lithuania, Volhynia, and northern 
Germany where it is now exterminated) ; in Asia south to the Altai. 
Still lives in northern Norway, Sweden, and Lapland." 

Norway. In former times it was common throughout the country, 
but only in the mountains in the southern part. At present it is 
rare in the high mountains and probably will soon become com- 
pletely extinct. Large bounties are paid for every specimen because 
of the damage the Wolverine does to cattle and reindeer. (Hj . Broch, 
in litt., December, 1936.) 

It is supposed that a small stock is left in the south of Norway 
on Hardangervidda and in Jotunheimen. While still found in the 
northern parts, it shows a considerable decrease there. It is ques- 
tionable whether it is possible to preserve the Wolverine in Norway. 
(Director of Forestry, Ministry of Agriculture, Norway, in litt., 
January, 1937.) 

Sweden. The Wolverine inhabits chiefly the mountain forests 
and the areas above the tree limit. It has therefore been mostly 
restricted to the country northward from northern Dalecarlia (lat. 
61 30' N.). In Wermland it was formerly found in small numbers, 
but disappeared before the middle of the last century. Single speci- 
mens have even been found as far south as Scania. At present, it 
occurs only from Jemtland northward. 

The skin of the Wolverine has been valued for centuries, and the 
animal has also been pursued because of its damage to livestock, 
mostly reindeer but also sheep to some extent. Consequently boun- 
ties are paid, and have recently been increased; the State pays 
10 Cr., while the Lappfund pays 100 Cr. for old animals and 50 Cr. 
for cubs. The total number of Wolverines killed in the whole 
country is, by decades, as follows: 

1856-1865 1,159 1896-1905 1,084 

1866-1875 1,201 1906-1915 717 

1876-1885 1,240 1916-1925 639 

1886-1895 992 1926-1934 517 

There is a pretty steady decrease in numbers from the beginning 
of the present century, indicating that there is danger of extermina- 
tion. (Einar Lonnberg, in litt., October, 1936.) 

Finland. Ognev (1935, p. 95) mentions the occurrence of the 
Wolverine about Lake Enara. 

Latvia. In the Baltic states the species was once found in large 
numbers, but now seems to have disappeared. In 1875 a specimen 
was killed near Gerki in Courland, and in 1876 another near Jacob- 
stadt. (Ognev, 1935, p. 94.) 

Lithuania. Its former existence is uncertain (T. Ivanauskas, 
in litt., November, 1936) . 



Poland. It was quite frequent in the eighteenth century and even 
in the nineteenth century, but is now most probably exterminated 
(M. Siedlecki, in litt., October, 1936). It lived till the end of the 
last century in Volhynia, Podolia, and Polesia (Lubicz v. Nieza- 
bitowski, 1934, p. 190) . 

Germany. Two reports of Wolverines in central or northern 
Germany in the eighteenth century are evidently based upon escaped 
captives (Blasius, 1857, p. 211; Hilzheimer, 1933, pp. 219-221). 

Russia. The species formerly ranged southward to the northern 
Ukraine. At present it is found rarely in the Western Area, and pos- 

FIG. 25. Wolverine (Gulo gulo) 

sibly in Volhynia. Once thought to have been exterminated in White 
Russia at the end of the nineteenth century, it may have survived 
till later. It formerly occurred in the Governments of Novgorod 
and Olonets. It ranges northward to the Kola Peninsula. Prior 
to 1901, 200-300 Wolverines were collected annually in the Govern- 
ment of Archangel, but at present there is a decrease. Prior to 1925, 
75 animals were captured annually in the Petchora district. The 
species now seems to be very rare in central Russia (Ivanovo Indus- 
trial Area and adjacent areas) . It is widely distributed in the Urals, 
south to about lat. 53 N. (Ognev, 1935, pp. 94-96.) 

Siberia. The range of the Wolverine extends from the Urals 
east to the Anadyr district and Kamchatka ; on the north it reaches 
the Arctic coast in places; on the south it extends to the Altai Moun- 
tains, the Tannu-Ola Mountains (Mongolia) , the Bargusin district, 
the Amur region, the Ussuri district (lat. 44 N.), and Sakhalin. 
Over this vast area its status varies considerably; it is reported as 
numerous in some places and as rare or absent in others: (Ognev, 
1935, pp. 97-100; map, p. 101.) 


The Wolverine is widely distributed in the forested regions of 
the U. S. S. R. as a whole (Russia and Siberia) . It is less common in 
northern Russia than in Siberia. Its decrease in central Russia 
results from the decrease of the forest cover. It is very common in 
parts of Siberia. It causes great damage to hunting interests, taking 
animals caught in traps, destroying hunters' provisions, attacking 
young ungulates and even adults in deep snow. It is not legally 
protected and may be killed at any time. (W. G. Heptner, in litt., 
December, 1936.) 

Sakhalin. "Schrenck and Kishida reported it from the island. It 
is ... a rare 'animal on Sakhalin." (Kuroda, 1928, p. 227.) A speci- 
men was taken in 1934 (Kuroda, 1938, p. 26) . 

Manchuria. "I heard sufficient from authentic sources to con- 
vince me that the animal is fairly common, at least in the northern 
forested area" (Sowerby, 1923, p. 71). 

Mongolia. The species is reported in the Tannu-Ola Mountains 
(Ognev, 1935, pp. 98, 100) . "I once saw a skin from the Urga district 
in Northern Mongolia" (Sowerby, 1923, p. 71). 

Economics. "Wolverine fur has been much in vogue of late 
years, and has consequently appreciated in value. For a good skin, 
thirty shillings is often asked." (Lydekker, 1901, p. 112.) 

The world's fur production for 1928 included 6,000 Wolverine 
skins [some probably from North America] (Jour. Soc. Preservation 
Fauna Empire, pt. 12, p. 64, 1930) . 

Folklore. Many curious bits of folklore concerning the Wolverine 
have been handed down. Some of them are quoted by Lloyd (1854, 
pp. 16-18) from Pontoppidan and Olaus Magnus. 

Family VIVERRIDAE: Civets, Mongooses, etc. 

This Old World family ranges over southern Europe, Africa, 
Madagascar, southern Asia, and the Malay Archipelago as far as 
Timor, Ceram, and the Philippines. There are about 40 genera and 
350-400 forms. Accounts of six forms are given here. 

Malay Binturong; Bear-cat; Black Marten. Bintoeroeng 



Viverra? Binturong Raffles, Trans. Linnean Soc. London, vol. 13, pt. 1, p. 253, 

1821. ("Malacca,") 
FIGS.: Geoffroy and Cuvier, Hist. Nat. Mammiferes, vol. 5, pis. 201, 202 

(subsp.?), 1824; Sclater and Sclater, 1899, p. 128, fig. 26; Lydekker, 1900, 

pi. 9, fig. 1 (subsp.?). 

Opinions differ as to the rarity of the Binturong, the six subspecies 
of which range from northeastern India and Tonkin through the 


Malay Peninsula to Sumatra, Java, Borneo, and Palawan. Never- 
theless, it is evidently in need of total protection, more especially 
because of the demands upon it by the Chinese for medicinal pur- 

There has long been uncertainty as to the exact taxonomic status 
of the various described forms, owing chiefly to the inadequacy of 
the series of museum specimens. The species as a whole is dis- 
tinguished from all other Viverridae by its long ear tufts and by 
the prehensile nature of its long, bushy tail. The fur is long, coarse, 
and black, more or less washed with gray, fulvous, or buff. (Lydek- 
ker, 1893-1894, p. 463.) The present subspecies is larger and darker 
than A. b. penicillatus; its winter coat is much shorter and less 
luxuriant with underwool than that of albijrons, and the long hairs 
are less extensively annulated with lighter color (Pocock, 1933, 
p. 1030). Body, 30 inches; tail, nearly the same (Raffles, 1821, 
p. 253). 

The Malay Binturong inhabits the Malay Peninsula north to 
Tenasserim and Siam, and also Sumatra. 

Siam. In this country the Binturong seems to be rare (Gylden- 
stolpe, 1919, p. 148) . Specimens have been recorded from Sikawtur, 
northwest of Raheng, western Siam; from Prachin, central Siam; 
from Sai Yoke, southwestern Siam; and from Bang Nara, Patani, 
Peninsular Siam (Kloss, 1917, p. 293, and 1919, p. 53; Gyldenstolpe, 
1919, p. 148). In Ratburi Province, southwestern Siam, "the 
Karangs are well acquainted with the animal and state it is 
generally distributed in evergreen forest" (Gairdner, 1915, p. 252). 
Since its habits are "largely nocturnal and arboreal, the Bear-cat 
is not easily obtained" (Kloss, 1917, p. 294) . 

Malay Peninsula. "The Bear-cat ... is generally obtained in 
Malacca, and is sometimes kept as a pet. It is easily domesticated, 
and becomes very affectionate, and will follow its master like a 
dog. It feeds on fruit, also taking small birds." (Ridley, 1895, p. 93.) 

"This delightful animal is apparently not uncommon on the main- 
land, but I have not heard of it occurring wild in Penang or Singa- 
pore. In the Museum at Taiping are specimens from Larut and 
Kuala Kangsar, Perak. It is represented in the Museum at Kuala 
Lumpor, and is said to be common in Selangor." (Flower, 1900, 
pp. 330-331.) 

In the Malay Peninsula "Arctictis, Hemigalus, Neofelis, . . . 
are not in my opinion vanishing forms. . . . The three carnivores 
are all rare but it is extremely difficult to estimate their status in a 
country covered with jungle. I see no reason why they should be 
classed under Vanishing forms' as there is plenty of country suited 
to their requirements and they are not systematically hunted. I 
prefer to regard them as uncommon animals, rarely collected. Never- 



theless, they are rare enough to be given total protection." (F. N. 
Chasen, in litt., March 31, 1937.) 

In Malaya "many with whom I talked . . . were insistent that 
such animals as the . . . binturong . . . are to-day practically 
non-existent. . . . 

"It is true that the loris and binturong fetch a high price in the 
Chinese market, but they are numerous." (Comyn-Platt, 1937t>, 
P- 48.) 

FIG. 26. Binturong (Arctictis binturong subsp.) 

Sumatra. In the Korinchi region two specimens are recorded 
from Sandaran Agong, 2,450 feet (Robinson and Kloss, 1918, p. 11). 

F. N. Chasen (in litt., May 5, 1937) considers the Binturong 
much more numerous in Sumatra than in the Malay Peninsula. 
"Most specimens seen in captivity originate on the east coast of 
Sumatra." It "needs protection as the Chinese use it for medicine." 

"In Sumatra, the binturong is found occasionally near Selat 
Pandjang. It is not often seen in Rokan and Bengkalis." (Heyn- 
sius-Viruly and Van Heurn, 1936, p. 63.) 

According to Dr. Hagen, the species is rare in Sumatra. This 
agrees with information obtained from natives inland from Palem- 
bang. During two years' residence in the Ogan Oeloe Subdivision 
only one animal was seen in captivity. (Coomans de Ruiter, 1932, 
p. 53.) 

Lyon (1908, p. 652) records specimens from Aru Bay, Sungei 
Mundau, Siak River, Pulo Payong, and Pulo Tebing Tinggi, eastern 


Sumatra. Pocock (1933, pp. 1018-1019) records additional speci- 
mens from Ulu If ok, Perak; Wellesley Province, Straits Settlements; 
Tenasserim Village; and Sanderan Agong, Sumatra. 

Nias Binturong 


Arctictis miasensis Lyon, Proc. U. S. Nat. Mus., vol. 52, p. 443, 1916. (Near 
Fadoro, Nias Island, off west coast of Sumatra.) 

Forty years ago this Binturong was "said not to be common" 
(W. L. Abbott, in Lyon, 1916, p. 443) . 

"Upper parts of back of head, neck, body, and all of tail, brownish 
black, coarsely and rather sparsely grizzled with ochraceous tawny 
on the lower back, sides, outerside of legs, and proximal two-thirds 
of tail; under parts tawny ochraceous." Tail, 540 mm. The ochra- 
ceous-tawny in the present form is replaced by buff or ochraceous- 
buff in A. b. binturong. (Lyon, 1916, p. 443.) Cranial and dental 
differences are discussed by Miller (1942, pp. 123-124). 

This Binturong is apparently restricted to Nias Island. Since the 
type specimen was described, a second individual has been recorded 
by Miller (1942, pp. 123-124) , who quotes the collector, Frederick A. 
Ulmer, Jr., as follows: "The young binturong was purchased alive 
from the natives of Soliga in Central Nias and was the only one I 
saw, although I heard of one other specimen in captivity near 
Gunong Sitoli." Ulmer also refers to it as a "rare animal." 

Banka Binturong 


Arctictis binturong kerkhoveni Sody, Natuurk. Tijdschr. Nederl. Indie, vol. 96, 
no. 1, p. 43, 1936. ("Banka Island," Malay Archipelago.) 

This form is based upon a tingle specimen from Banka Island. 

It is the smallest subspecies. The fur is black, with short buffy 
tips to the hairs in some places, especially the head and forelegs; 
tail plain black, with light buffy bases to hairs on basal two-thirds 
of the ventral surface. Head and body, 600 mm.; tail, 520 mm. 
(Sody, 1936, p. 43.) 

No information is at hand concerning the frequency of the Bin- 
turong on Banka. 

Himalayan Binturong 


Paradoxurus albifrons F. Cuvier, Mem. Mus. Hist. Nat. [Parisl, vol. 9, p. 48, 
pi. 4, upper fig., 1822. (Based upon a menagerie specimen in Bengal, 
said to have come from Bhutan.) 


FIGS.: Wolf, 1867, pi. 10; Royal Nat. Hist., vol. 1, p. 463, 1893-94; Jour. 
Bombay Soc. Nat. Hist., vol. 38, no. 2, suppl., pi. 60, 1935. 

This form ranges through Nepal, Sikkim, Bhutan, Assam, Upper 
Burma, and Tonkin. 

It is distinguished from other forms by the length of its winter 
coat and the abundance of long underwool; the color of the body 
varies from jet black to tawny'or gray (Pocock, 1933, p. 1030). 

"The Binturong appears to be rare in Northern India." Speci- 
mens are recorded from Assam; from Endwagyi Lake, in Myitkyina, 
Upper Burma; from Fouine, Tonkin; and from Lower Laos. Only 
one skull appears to be known. (Pocock, 1933, pp. 1016-1017, 1030.) 

"Mr. W. L. Sclater says that it is found even as far west as Simla" 
(Lydekker, 1900, p. 334). 

Javanese Binturong 


(Original reference not found; not in Sherborn; cf. Temminck, Monographies 
Mammalogie, vol. 2, p. 310, 1841. Possibly the name was first published by 
Temminck sometime prior to 1825 in the prospectus of his "Monographies" 
(cf. Valenciennes, 1825, p. 57, footnote). Pocock's citation (1933, p. 1031), 
"Temminck, Mon. Mamm. ii. p. 18, 1835," is evidently erroneous; he 
gives "Java" as the type locality.) 

FIGS.: Ann. Sci. Nat., vol. 4, pi. 1, 1825; Coomans de Ruiter, 1932, p. 54, fig. 12. 

The range of this form is Java and Borneo. 

It is distinguished from A. b. binturong by its smaller skull and 
generally lighter color; pale annulation of the hairs extensive and 
profuse (Pocock, 1933, p. 1031). Head and body, 960 mm.; tail, 
890 mm. (Schwarz, 1911, p. 636) . 

Java. Temminck (1841, vol. 2, p. 311) was inclined to consider 
the Binturong the rarest of the mammals of Java and Sumatra. The 
species is "apparently rare in Java" (Shortridge, in Thomas and 
Wroughton, 1909, p. 386). 

Heynsius-Viruly and Van Heurn write (1936, p. 63) : 

Few data were received concerning this animal. It was observed in Java 
in the vicinity of Madjalengka and near the border of Tomo .... 

The binturong is often seen in the Midangan mountains, but it is much 
scarcer on the Andjasmoro. None were seen there for the past two years. . . . 

Owing to his size, his striking color, his pretty fur and his rather sluggish 
motions, the binturong is doomed to be soon exterminated, unless very stern 
measures are taken. It occurs on all the Greater Sunda Islands, but is nowhere 
common. . . . Young animals become very tame and affectionate in captivity. 

This pretty animal . . . should be intensively protected. 

Borneo. "These animals are common in parts of Borneo, usually 
living in the dense forest, but when in search of fruit they will often 
visit gardens" (Hose, 1893, p. 24) . 


Schwarz (1911, p. 636) mentions specimens from Sandakan and 
La Datu, North Borneo, and from Sarawak. Pocock (1933, p. 1031) 
records others from Mount Mulu, Mount Dulit, and Saribas in 
northern Borneo. In the Western Division of Borneo the Binturong 
is not rare, and specimens are frequently kept in captivity (Coomans 
de Ruiter, 1932, p. 54). 

Palawan Binturong 


Arctitis [sic] whitei J. A. Allen, Bull. Am. Mus. Nat. Hist., vol. 28, p. 15, 
1910. ("Iwahig, Palawan, Philippine Islands") 

This form is apparently restricted to Palawan. 

It is closely related to A. b. penicillatus, but has a smaller skull; 
general color black, strongly washed with fulvous; hairs of nose 
and facial region tipped with whitish; neck all round and terminal 
fifth of tail black; ears narrowly bordered with white or yellowish. 
Head and body, 700 mm.; tail, 610 mm. (J. A. Allen, 1910, p. 15.) 

Only about four specimens of this form seem to be on record 
(Taylor, 1934, p. 357). 

Fossane; Lesser Fossa 

FOSSA FOSSA (Schreber) 

Viverra Fossa Schreber, Saugthiere, vol. 3, pi. 114, 1776, and p. 424, 1777. 

(Based upon "la Fossane" of Buffon (Hist. Nat., vol. 13, p. 163, pi. 20, 

1765); type locality, "Madagascar.") 
SYNONYM: Fossa daubentonii Gray (1865). 
FIGS.: Buffon, op. tit., pi. 20; Schreber, op. cit., pi. 114; J. E. Gray, 1873, pi. 74. 

This endemic viverrid of Madagascar is accorded special pro- 
tection as a Class A species under the London Convention of 1933. 

The ground color is light ashy gray, slightly washed with red- 
dish; stripes and spots blackish brown; breast, belly, and legs gray; 
tail gray, incompletely ringed with brown (Schreber, 1777, p. 424). 
Gray (1873, p. 872) gives the following description: "Brown or red- 
dish, closely grizzled with an abundance of white hairs, with four 
rows of more or less confluent black spots on each side of the 
back, a few black spots on the hinder thighs. The chin, neck, and 
belly whitish, more or less obscurely spotted." Daubenton (in 
Buffon, 1765, p. 166) gives the following measurements: head and 
body, 17 inches; tail, 8J inches. 

Our information concerning this animal is very meager. Accord- 
ing to Buffon (1765, p. 164), in captivity it eats flesh and fruit and 
is especially fond of bananas. 

The Mission Zoologique Franco-Anglo-Americaine of 1929-1931 
obtained 13 specimens (Delacour, 1932, p. 220). 


"The striped civet of the humid forest is apparently entirely 
nocturnal .... Though fairly common" 20 kilometers west of 
Vondrozo, "as we found by trapping, none was seen in the daytime. 

"Three stomachs from near Vondrozo and one from near Maro- 
antsetra all contained insect matter and one contained also a lizard 
.... The striped civet was known as 'fanaloka' amongst the Ata- 
moor in the southeast." (Rand, 1935, p. 93.) 

Family PROTELIDAE: Aard-wolves 

The single genus of this family contains one species, which has 
been divided into half a dozen subspecies. They range over southern 
and eastern Africa. All come within the scope of this work. 

Aard-wolf. Maanhaar Jackal (Boer). Faux-loup; Loup de 
terre (Fr.). Zibethyane (Ger.) 


Viverra cristata Sparrman, Resa till Goda Hopps-Udden, vol. 1, p. 581, 1783. 

(Near Little Fish River, Somerset East, Cape Province.) 
FIGS.: Cuvier, Regne animal, disciples' ed., Mamm., atlas, pi. 40, fig. 3, 

1836-1849; Proc. Zool. Soc. London 1869, pi. 36; W. L. Sclater, 1900, 

vol. 1, p. 81, fig. 21; Anderson and de Winton, 1902, pi. 28; Derscheid, 

1925, pi. A; Pocock, 1937, p. 758, fig. 

The Aard-wolf is of particular scientific interest as the sole repre- 
sentative of the family Protelidae; and it has been accorded rigid 
protection as a Class A mammal under the London Convention of 

Six subspecies have been proposed, as listed below, but since their 
distributional limits have not been worked out, all will be included 
in this account of the species as a whole. 

Proteles cristatus cristatus (Sparrman) . Cape Aard-wolf. (Type 
locality as given above.) 

Form hyenalike; general color dirty yellowish gray, with project- 
ing coarse hairs, black and white; an erectile black mane from 
nape to tail; seven to nine transverse black stripes on sides; upper 
parts of limbs with indistinct black bands; feet black; face, lower 
jaws, and chin brown; tail yellowish at base, rest black. Head and 
body, 32 inches; tail without hairs, 6 inches. (W. L. Sclater, 1900, 
vol. 1, pp. 80-81.) 

Proteles cristatus pallidior Cabrera, Ann. Mag. Nat. Hist., ser. 8, 
vol. 6, p. 464, 1910. Nubian Aard-wolf. ("Suakim," Anglo-Egyptian 

General color pale yellowish cream ; body hairs unicolored ; cheeks 
and sides of neck not rufous as in true cristatus; mane with very 
little black; feet brownish, not black; tail black only at tip (Cabrera, 
1910, p. 464). 


Proteles cristatus septentrionalis Rothschild, Novit. Zool., vol. 9, 
p. 443, 1902. Somali Aard-wolf. ("Somaliland.") 

Creamy white, washed with buff on neck and sides of rump; 
stripes less defined than in cristatus; mane black, variegated with 
creamy white (W. Rothschild, 1902, p. 443). 

Proteles cristatus termes Heller, Smithsonian Misc. Coll., vol. 61, 
no. 13, p. 9, 1913. Masailand Aard-wolf. ("Headwaters of the 
Amala River west of the Loita Plains, British East Africa.") 

Interorbital region black, crown grizzled, feet black, and ears 
blackish as in cristatus; body stripes narrow; ground color grayish 
fulvous; tail black on terminal third; muzzle and chin black; throat 
grayish buffy. Head and body, 680 mm.; tail, 310 mm. (E. Heller, 
19136, p. 9). 

Proteles cristatus transvaalensis Roberts, Ann. Transvaal Mus., 
vol. 15, pt. 1, p. 6, 1932. Transvaal Aard-wolf. ("Roodekuil, Pre- 
toria," Transvaal.) 

Pale buffy; face, bands on limbs and body, and distal third of 
tail dark brown; mane rufous-white, the hairs with three brown 
bands and blackish tips. Head and body, 650 mm.; tail,, 270 mm. 
(Roberts, 1932, p. 6.) 

Proteles cristatus harrisoni Rothschild, Novit. Zool., vol. 9, p. 443, 
1902. Angola Aard-wolf. ("Umpata, Mossamedes district, S. An- 

Head white, grizzled with black; body pale orange rufous; stripes 
less developed than in other races; mane and tail black, variegated 
with rufous (Rothschild, 1902, p. 443) . 

The species as a whole has a wide range over southern and eastern 
Africa, from Cape Province and Natal north to Angola, Northern 
Rhodesia, and Anglo-Egyptian Sudan. It is apparently absent 
along the low-lying east coast from the Transkei district of Cape 
Province to Portuguese East Africa. (Shortridge, 1934, vol. 1, 
p. 150.) 

Anglo-Egyptian Sudan. Specimens have been recorded from the 
vicinity of Suakin (Anderson and de Winton, 1902, p. 198; Cabrera, 
1910, p. 465) and from the Blue Nile (Butler, in Maydon, 1932, 
p. 151). 

Eritrea. This animal is rather widespread, if not common, in the 
greater part of the country, especially in the southwest (Zammarano, 
1930, p. 77). 

Ethiopia. It is "common on the lowlands and foot hills wherever 
white ants abound" (Wylde, 1901, p. 485). 

British Somaliland. "The aardwolf is found sparsely scattered 
throughout Somaliland. They are almost invariably seen alone." 
(Drake-Brockman, 1910, p. 38.) 


Italian Somaliland. Recorded by De Beaux (1935, p. 12). 

Kenya. Hollister (1918, pt. 1, p. 138) lists specimens from the 
following localities: Kabalolot Hill, Sotik; Northern Guaso Nyiro 
River; Telek River, Sotik; and Ulukenia Hills. 

Tanganyika Territory. Recorded from Tabora (Matschie, 1895, 
p. 62). 

Northern Rhodesia. "I am reliably informed that specimens 
have been obtained at Tara and Kalomo in the Batoka Province. 
Elsewhere the natives do not seem to have heard of it." (Pitman, 
1934, p. 162.) 

Southern Rhodesia. It ranges rather sparsely over this country 
(western Matabeleland, etc.) (Shortridge, 1934, vol. 1, p. 150). 

Transvaal. "In the Eastern Transvaal the Aard Wolf is not 
found in the low-veld proper; it occurs in the more open country 
among the foothills of the Drakensberg at a height of over 1,500 
feet" (Hamilton, in Shortrklge, 1934, vol. 1, p. 150). A specimen is 
recorded from Potchefstrom (W. L. Sclater, 1900, vol. 1, p. 82). 

Natal. "It is fairly common in Natal" (Warren, in Shortridge, 

1934, vol. 1, p. 150). 

Cape Province. The Aard-wolf is reported as not uncommon 
throughout the colony (W. L. Sclater, 1900, vol. 1, p. 81). 

Bechuanaland. "The Kalahari Sand-Plains" are "perhaps the 
regions in which it is most plentiful. . . . The karross-making 
tribes in Bechuanaland . . . are said to procure most of their aard 
wolf skins with the aid of dogs." (Shortridge, 1934, vol. 1, p. 150.) 

As many as 14 have been seen together in the Kalahari (Langdon, 
in Shortridge, 1934, vol. 1, p. 151). 

South-West Africa. "Proteles is widely distributed throughout 
South -West Africa; nowhere very abundantly. It is apparently 
rather scarce along the valley of the Orange River, and northwards 
in the neighbourhood of the Okavango and in the Caprivi. . . . 

"The Aard Wolf is fairly plentiful around Gobabis and in the 
sand-plains generally; and is also familiar in Namaqualand, Da- 
maraland, the Kaokoveld, Ovamboland, and the Namutoni Game 
Reserve." (Shortridge, 1934, vol. 1, pp. 149-150.) 

Angola. It is rather common in the south of Angola but much 
rarer in the north. A skin was brought in to Vila da Ponte, where 
the animal was unknown to the natives. (Monard, 1931, p. 66, and 

1935, p. 228.) The type locality of the subspecies harrisoni is Um- 
pata in the Mossamedes district. 

Economic status. "The coat is very handsome, and ... its skin 
is more sought after [than that of the hyenas] by some of the native 
tribes notably the Bechuanas, who hunt and trap it systematically" 
(Bryden, 1899, p. 599). 

"Sparrman and other authors who have examined the stomachs 



of these animals, found that they contain nothing but termites or 
white ants; this is further confirmed by Mr. Cloete, who writes that 
he has examined the stomachs of more than fifty, and never found 
any trace of anything else than a purely insectivorous diet, ants 
being the chief constituent." However, farmers report that this 
species kills kids and lambs merely for the sake of the milk con- 
tained in their stomachs. (W. L. Sclater, 1900, vol. 1, p. 82.) 

The food consists of insects particularly termites, locusts, beetles, 
and grubs. The animal has been accused of killing lambs and kids, 

Fie. 27. Aard-wolf (Proteles cristatus subsp.) 

but evidently without justification and through confusion with the 
Jackal. Its weak dentition is sufficient evidence of its harmlessness 
in respect to livestock. Yet, despite its inoffensive nature and the 
distinct service it performs in destroying such agricultural pests as 
termites, it was, in former years, officially listed by several govern- 
ments as "harmful," and a bounty of half a pound sterling was paid 
in the Cape Province for each Aard-wolf killed. (Derscheid, 1925, 
P. [78].) 

It feeds to some extent on "small rodents, reptiles, and the nest- 
lings and eggs of ground-nesting birds" (Shortridge, 1934, vol. 1, 
p. 151). 

In South Africa generally it is subjected to a great deal of irre- 
sponsible persecution, and is becoming scarce in farming and other 
settled districts. However, it is in no immediate danger of actual 
extermination. (G. C. Shortridge, in litt., October 14, 1937.) 


Although normally the animal has no unpleasant odor, it is able 
to eject an evil-smelling fluid from its anal glands as a defense 
against such enemies as dogs. 

Family FELIDAE: Cats 

This family is nearly cosmopolitan, but it does not occur natur- 
ally in Greenland, the eastern Malay Archipelago, or Australasia. 
Twenty or more genera are recognized by some authorities, and 
there are probably more than 250 forms. Dr. Allen discusses 12 
North American forms in the preceding volume (1942), while 27 
Old World forms are dealt with in the following pages. Man's 
prejudice against some of the larger members of the cat family 
(such as Lions, Tigers, Leopards, and Cougars) is linked with his 
necessary efforts to defend himself or his livestock from their attacks. 



Cryptoprocta jerox Bennett, Proc. Zool. Soc. London 1833, p. 46, 1833. ("Mada- 

FIGS.: Bennett, 1834, pi. 21; Schreber, Saugthiere, suppl. vol. 2, pi. 125CC, 
1841; Schlegel and Pollen, 1868, pi. 8; Proc. Zool. Soc. London 1895, pi. 
26; Beddard, 1902, p. 405, fig. 199; Elliot, 1907, p. 397, fig. 43; Kaudern, 
1915, pi. 3, fig. 2; Sibree, 1915, pi. facing p. 302; Pocock, 1937, p. 760, fig. 

This largest carnivore of Madagascar occurs in limited numbers 
and will probably require protection if it is to survive in the 
dwindling forests of that great island. It is an endemic species. 

Since Bennett's type specimen was distinctly immature, the fol- 
lowing description of an adult male is derived from Schlegel and 
Pollen (1868, pp. 13-14). Hairs of upper parts ringed with brown 
and pale reddish yellow; lower parts of head and body uniform 
reddish yellow, taking on a strong rusty tint toward the middle of 
the venter. Total length, 56 inches; tail, 26 inches. 

Bennett (1834, p. 140) quotes Charles Telfair as follows: "It is 
the most savage creature of its size I ever met with: its motions 
and power and activity were those of a tiger: and it had the 
same appetites for blood and destruction of animal life." 

Milne Edwards and Grandidier write (1867, p. 317) that it occurs 
rather commonly on the west coast, from the River Mangouke 
[Mangoky] northward. Three specimens were secured between 
Morondava and Manharrive [Maharivo?]. The animal often carries 
off goats and especially kids. 

This animal is very carnivorous and is endowed with great 
strength. It is dangerous to man only when wounded or in rut. 
At other times "it steadily flees from man. At the mating season 


it is often seen in bands of four to eight individuals. It is said to be 
fond of lemurs and to pursue them in trees. It is also destructive 
to poultry, young pigs, and other domestic animals. The natives 
really fear this species, but they enjoy its flesh. (Schlegel and 
Pollen, 1868, pp. 15-16.) 

According to Milne Edwards and Grandidier (1875a, p. 341, foot- 
note) , this is the only native animal of Madagascar that the Saka- 
lava (a western tribe) have been able sometimes, but very rarely, 
to train for hunting the Wild Hog (Potamochoerus larvatus) . 

Kaudern states (1915, pp. 79-80) that Cryptoprocta appears to 
be distributed over the entire island and that it is probably nowhere 

FIG. 28. Fossa (Cryptoprocta jerox). After photo in Brehm. 

rare. In northwestern Madagascar it was very common. He saw 
the animal three times in the wild at Ste. Marie de Marovoay on 
the Betsiboka River, and its tracks were observed everywhere in 
the sand. One was killed there in a poultry yard. Another was 
secured at Katsepe on the Bay of Bombetoke, and two live young 
ones were brought in by natives at Andranolava, in north central 
Madagascar. Black individuals are reported from the interior and 
from the great rain forests on the east coast. 

According to Sibree (1915, pp. 302-303), the northwest coast is 
the animal's "special habitat. This creature is called by the people, 
Fosa . . . , and although small is very ferocious .... Examples 
of the fosa have been seen in the outskirts of the upper belt of forest 
on the east side of the island .... A specimen I once saw was of a 
beautiful black colour, but I believe this was only a variety, and 
not a distinct species from the brown animal. The fosa is much 
dreaded by the Malagasy, and, from its mode of attack, appears to 
be like an immense weasel, attacking large animals, such as the 
wild boar and even oxen." 

G. M. Allen (1918, p. 514) records a specimen from the vicinity 
of Tulear. 


Petit (1931, p. 588) records a female and its three young ones 
captured in 1922 in the region of Tamatave, and two young ones 
taken in the region of Fenerive on the east coast. 

The Mission Zoologique Franco-Anglo-Americaine of 1929-1931 
secured 6 specimens of this species, as compared with 13 specimens 
of Fossa jossa (Delacour, 1932, p. 220). 

Rand (1935, pp. 93-94) says: 

The fossa inhabits the rain forest of the east and the dryer forest of the 
west at least as far south as Tabiky [inland from Cape St. Vincent], and was 
well known to the natives. [Two were seen near Tsarakibany and Maromandia 
during the daytime, though the natives said it was nocturnal.] This viverrid 
was much disliked by the natives because of its raids on their fowls. Twice 
I saw fossa skins in the possession of natives, but this was probably due to 
European influence as the natives rarely use mammal skins for any purpose. 
One large fossa was brought to me that had been run down with dogs and 
speared. From the natives we heard no accounts of its attacking sheep or 
young cattle and its reputation in literature for ferocity and the fear with 
which it is regarded by the natives is exaggeration. My gun boy had a 
particular antipathy for it because, he said, in his country near Vondrozo, 
where the dead are walled up in caves, the fossa sometimes dug out the 
corpses and fed on them. The natives universally called it "fossa." 

European Wildcat. Chat sauvage (Fr.). Wildkatze (Ger.). 
Gato monies; Gato salvage (Sp.) Gatto selvatico (It.) 


Felis (Catus) silvestris Schreber, Saugthiere, vol. 3, p. 397, pis. 107A, 107 Aa, 
1777. (Germany.) 

FIGS.: Gervais, Hist. Nat. Mammif., pt. 2, pi. 17, 1855; Blasius, 1857, p. 162, 
fig. 101; Elliot, 1883, pi. 30; Royal Nat. Hist., vol. 1, pi. facing p. 422, 
1893-94; Hamilton, 1896, frontisp.; Martin, 1910, pi. 27; Cabrera, 1914, 
pi. 9, fig. 1; Zeitschr. f. Saugetierk., vol. 7, pi. 7, fig. 7, 1932; Colosi, 1933, 
p. 55, fig.; Didier and Rode, 1935, p. 283, fig. 163; Schmidt, 1938, pi. 5. 

The typical European Wildcat shows a very general and marked 
recession in France and central Europe, amounting to extirpation 
in many parts of its former range. Apparently its chief remaining 
stronghold is in the Balkan countries. 

More definite light is needed on the question as to whether inter- 
breeding with feral Domestic Cats takes place at all or on a 
sufficient scale to menace the Wildcat's survival as a pure-bred 
species. Fatio states (1869, p. 276) that hybrids are sometimes 
met with in Switzerland, and that he has examined a number of 
specimens; the pelage, he adds, is often spotted with white. Ferrant 
(1931, p. 62), in discussing the Wildcat in Luxembourg, says that it 
mates frequently with feral Domestic Cats. Prof. M. Hirtz refers 
(in litt., December, 1936) to hybrids in Yugoslavia, and the National 
Council for Nature Protection does likewise (in litt., October, 1936) 
in Poland. On the other hand, Pocock (1907, pp. 165-166) is rather 
skeptical in regard to the alleged interbreeding. 


Hamilton speaks rather emphatically in his monograph (1896, 
p. iv) : "On a careful examination of a number of examples of the 
Wild Cat of the present time I found many indications of a mixture 
of the two races. 

"It would seem as if the original Wild Cat, as it existed in the 
olden days, has been almost exterminated throughout Europe, and 
that its place has been taken by a mongrel race, the result of con- 
tinual interbreeding during many centuries (2000 years) of the 
Wild and the imported Domestic Cat." 

The Wildcat is slightly larger than the Domestic Cat; fur longer, 
this being especially noticeable in the tail ; general color approaching 
the smoke gray of Ridgway; dark markings on sides and legs tend- 
ing to be faint, brownish, and ill-defined; tail, abruptly rounded 
at the black tip, with two to four more or less complete black rings ; 
tip of ear slightly blackish. Head and body, 481-545 mm.; tail, 
309-310 mm. (Miller, 1912, pp. 457-463.) Head and body, 450- 
700 mm.; tail, 200-300 mm. (Didier and Rode, 1935, p. 284). 

The typical subspecies ranges from northern Spain, France, and 
northern Germany eastward to Poland and Russia, and southward 
to Italy and the Balkan Peninsula. 

In Spain it inhabits the Pyrenean, Cantabrian, and north central 
districts, and it may extend to northern Portugal. Probably the 
Douro and the Ebro constitute its southern limits. (Cabrera, 1914, 
p. 204.) "Curiously enough there is no specimen in the Museums 
of Ponferrada, Lugo, and Santiago. I was left in doubt as to whether 
the wild cat occurs in the Sierra de Picos, in Galicia, or in the Picos 
de Europa." (Gadow, 1897, p. 367.) 

In France, where it was formerly very common, the Wildcat is 
at present in the process of disappearing, but may still be met with 
in certain regions, such as the Ardennes, the Forest of Orleans, the 
Pyrenees-Orientales, and the Forest of Carnelle. It preys upon 
rabbits, hares, squirrels, rats, young Roebucks and Wild Boars, 
grouse, partridges, and pheasants, and even eats fishes. It is a very 
injurious animal, which one need not hesitate to destroy. (Didier 
and Rode, 1935, p. 287.) 

Hunting has contributed in part to its depletion in France, but 
does not explain this altogether. The species is very much in danger, 
and we do not think that protective measures can be effective. (E. 
Bourdelle, in litt., March 6, 1937.) 

In Belgium the species has become very rare, but still exists in 
some forests in the Province of Luxembourg. It is systematically 
destroyed as an injurious animal, while another cause of depletion 
is deforestation. Total protection ought to be adopted. (Musee 
Royale d'Histoire Naturelle de Belgique, in litt., September, 1936.) 


In the Grand Duchy of Luxembourg the Wildcat is rather com- 
mon in the extensive forests of the Ardennes, in the environs of 
Echternach, Grevenmacher, Manternach, and Fischbach, in the 
Grunewald, etc. (Ferrant, 1931, p. 61). 

In The Netherlands the species was long since exterminated (Van 
den Brink, 1931, p. 174). 

In Germany the Wildcat has survived better than the larger 
carnivores; it occurs in very small numbers in the Bavarian moun- 
tains, the Black Forest, the Odenwald, and the Riesengebirge. From 
1850 to 1860 ten animals were killed in Gotha; in 1885-86, two in 
Silesia; in 1928 an unquestionably pure-blooded male was taken in 
the Harz Mountains, and in the same year a male in the Kurische 
Nehrung. (Krumbiegel, 1930, pp. 5-6.) The Wildcat is still regu- 
larly observed in the Eifel, in the Moselle Mountains, and in the 
Hunsriick, and there is one from the Pfalz in the Koln Zoological 
Garden (Hauchecorne, Zeitschr. f. Saugetierk., vol. 9, p. 4, 1934). 
The animal is almost exterminated in Germany, and is protected 
as a natural monument (Internationale Gesellschaft zur Erhaltung 
des Wisents, in litt., October, 1936) . 

In Denmark bones of the Wildcat have been found in kitchen 
middens, but there is no record within historical times (Winge, 
1908, p. 116). 

In Switzerland it appears to have been abundant in the sixteenth 
century and was then the object of much hunting; but it had 
become rare by the beginning of the nineteenth century. In the 
1860's some were killed each year in the Alps and the Jura, and a 
few were still found in the cantons of Bern, Lucerne, Unterwalden, 
Uri, Schwyz, Glarus, Thiirgau, and Valais. The most were found in 
the Jura region, from Geneva to Basle. The species seemed to have 
disappeared from Ticino. (Fatio, 1869, p. 275.) During the last 
decades it has become very rare and is probably extinct, although 
it is possible that a few survive in the forests of the Alps and the 
western Jura (Federal Forest, Game, and Fish Inspection, Bern, 
in- litt., March, 1937) . 

Wildcats have almost disappeared from northern Italy, and are 
rare everywhere except in the Maremma, in the southern provinces 
of Gargano, and in Calabria (Colosi, 1933, p. 56). [The animal of 
the Tuscan Maremma is regarded by Martorelli (1896, p. 266) as 
identical with the Sardinian Wildcat (Felis sarda).] The Wildcat 
still occurs in the Sila Mountains of Calabria (Hecht, 1932, p. 23). 
According to the Laboratorio di Zoologia Applicata a Caccia (in 
litt., September, 1936) , the animal is scattered through Sicily as well 
as the Italian Peninsula ; in legislation it is rated as a harmful species. 

In former times it was probably found everywhere in Austria. 
It is now exterminated in Burgenland but is said to survive in 


Rosenbachtal in Carinthia. In 1926 two specimens were killed in 
the district of Volkermarkt, Carinthia. In Lower Austria the last 
one was killed in 1912. The species is no longer found in Salzburg. 
In the Tyrol 26 specimens are said to have been shot in 1876. By 
1888 it was almost exterminated in northern Tyrol, but was con- 
sidered more frequent in southern Tyrol. In Vorarlberg it was 
reported as late as 1918, but has now disappeared. (G. Schlesinger, 
in Hit., March, 1937.) 

Within the boundaries of the present Hungary it was common 
before World War I, but is now decreasing. It has no legal pro- 
tection. It is also found in the northern part of the former Hungary 
(now Czechoslovakia) , but not together with the Lynx. ( J. Schenk, 
in Hit., November, 1936.) 

In comparison with its status in most other parts of Europe, 
the Wildcat is comparatively common in Yugoslavia. Considerable 
numbers are found only in certain regions, and especially in the 
enormous oak forests of Slavonia. The statistics are uncertain, 
since they probably include hybrids and feral Domestic Cats. The 
reported annual kill from 1891 to 1921 was about 500 to 1,000 
specimens, reaching a maximum of 1,207 in 1904. The minimum 
kills were 420 in 1918 and 331 in 1931. These figures pertain chiefly 
to Croatia and Slavonia. (M. Hirtz, in litt., December, 1936.) In 
northwestern Croatia the animal is rare near Jasenak and near 
Otocac (Wettstein, 1928, p. 35). 

Lord Lilford wrote of frequently meeting with Wildcats in the 
Province of Epirus, near the boundary between Albania and Greece 
(Hamilton, 1896, p. 35). 

The Wildcat is found throughout Greece except on the islands. 
A decrease has been observed, but there is no danger of extermina- 
tion. (Game Department, Ministry of Agriculture, Greece, in litt., 
October, 1936.) (The Wildcat of the Peloponnesus is discussed on 
a subsequent page, under the name of Felis silvestris morea.) 

Turkey is included in the range by Blasius (1857, p. 166). 

In Bulgaria the Wildcat occurs in considerable numbers and is 
in no danger of extermination. The skins are marketed locally. 
(Bulgarian Game Association "Sokol," in litt., February, 1937.) 
H. W. Shoemaker (in litt., June 30, 1932) is of the opinion that 
the Wildcats in this country are being rapidly destroyed. 

The species is common in Rumania, occurring in nearly all forests 
of greater or lesser altitude; it is also frequently found in the 
lowland forests (District of Ilfoo) and in the flood lands of the 
Danube (Calinescu, 1930, p. 366). 

In Poland it occurred formerly from the Carpathians to the 
Baltic but is now met with chiefly in the eastern Carpathians; it 
is also found in the forests along the Dniester (Zurawno, Stanis- 


lawow) and in Podolia. It is nowhere common (Niezabitowski, 
1934, pp. 190-191). By 1936 the species was considered restricted 
to the Carpathians, where its numbers are roughly estimated at 300. 
It is protected from February 1 to September 30. (M. Siedlecki 
and National Council for Nature Protection, Poland, in litt., October, 
1936.) The proposed International Tatra Park and the proposed 
International Park of Poland, Czechoslovakia, and Rumania will, it 
is hoped, provide absolute protection for the Wildcat and other 
species (Benedyct Fulinski, MS., 1933). 

The following summary of the Wildcat's status in Russia is de- 
rived from Ognev (Ogneff, 1930, pp. 55-58) . The information is frag- 
mentary and suffers from lack of material. Pallas (1811-1831) de- 
nied the animal's existence throughout Russia except in the Caucasus. 
Georgi (1800) reported it in the southwestern governments, on the 
Dniester, and in the central Urals (Bashkiri). Brandt (1853) be- 
lieved in an early, much wider distribution, as far as the central gov- 
ernments and perhaps to the Urals. Kessler (1856, 1858) records the 
Wildcat in Volhynia and Podolia. In 1854 it was reported in the 
Governments of Grodno, Vitebsk, and Kovno. Sabaneeff (1878) 
considered the Government of Minsk the center of its distribution 
in western Russia. According to Charlemagne (1920), it is now 
very rare in Volhynia and in the vicinity of Odessa and Tiraspol 
(Government of Kherson). Old reports from central and northern 
Russia in the latter part of the past century may have been based 
upon feral Domestic Cats. The question as to whether the Wildcat 
ever occurred in the Urals is unsolved. 

On the subject of general depletion Elliot comments (1883, text 
to pi. 30) : "Various are the causes that have effected this ; probably 
the chief one is the constant persecution to which the animal has 
been subjected, as this species has but few friends, and no quarter 
is shown when it is met with in the forest." 

Hamilton (1896, pp. 31, 95) remarks on the spread of the Domestic 
Cat with the increase of the human population, and the resulting 
interbreeding with the Wildcat, as possibly a chief factor in the 
disappearance of the pure-bred wild animal in Europe generally. 

British Wildcat 


Felis grampia Miller, Ann. Mag. Nat. Hist., ser. 7, vol. 20, p. 396, 1907. 

("Invermoriston District, Inverness, Scotland.") 
Fios.: Millais, 1904, pis. facing pp. 166, 170, 172, 174, 178; Pocock, 1937, 

p. 777, fig. 

The British Wildcat formerly occurred throughout Great Britain 
but is now restricted to the wilder portions of Scotland (Miller, 
1912, p. 464). 


It is like Felis silvestris silvestris of continental Europe, but the 
general color is darker, approaching broccoli-brown; dark markings 
on sides and legs tending to be extensive, blackish, and well defined ; 
upper side of feet and inner surface of hind legs ochraceous-buff, 
under side of body duller; intercrural and pectoral white areas well 
defined; middle of chest mottled with black; dark markings on tail, 
legs, and upper parts similar to, but more definite than, those of 
F. s. silvestris (Miller, 1912, p. 464). Males: head and body, 558- 
660 mm. ; tail, 280-355 mm. Females slightly smaller. (Millais, 1904, 
p. 170.) 

The following account is condensed from Millais (1904, pp. 170- 
180) . It is not known when the Wildcat became extinct in southern 
and central England, but it probably lingered until the forests were 
cleared. In Wales it may have survived till about the end of the 
nineteenth century. Approximate dates of last records in England 
are: Yorkshire, 1840; Lake District, 1843. In churchwardens' ac- 
counts and other records there is mention of bounties paid for Wild- 
cats in the sixteenth, seventeenth, eighteenth, and even nineteenth 

Last dates in certain counties of Scotland are: Berwickshire, 
1849; Dumfriesshire, Wigtown, and Kirkcudbright, about 1832; 
Dumbarton, 1857; Perthshire, 1870-71; Aberdeenshire, 1891; Forfar, 
Kincardine, Banff, Elgin, and Nairn, practically extinct since 1850. 
"Northern and western Inverness is, with western Ross-shire, the 
main stronghold of the Wild Cat to-day." Up to 1904 William 
Macleay, of Inverness, annually received eight or ten specimens, 
chiefly from Glenmoriston and Balmacaan, west of Loch Ness. "In 
Sutherland the Duke of Sutherland does not allow the slaughter of 
Wild Cats to take place," and the animals are on the increase in 
certain parts. In Caithness the species was never common but was 
reported as occasional until 1845, and it evidently survived to a 
somewhat later date. 

The Wildcat's prey includes poultry, lambs, and roe fawns. But 
since it keeps down certain animals such as grouse, hares, and 
rabbits that are considered undesirable in deer forests, it receives 
protection from the sportsmen who control these forests. 

According to the minutes of a meeting of the Society for the 
Preservation of the Fauna of the Empire in October, 1922, "a cir- 
cular to owners and tenants of deer forests in Scotland, asking them 
to protect wild cats and martens, had been well received." Several 
years later it is reported that "we get very gratifying reports in 
regard to the preservation by land owners of wild cats and pole cats" 
(Onslow, 1929, p. 7). 

"Mr. N. B. Kinnear remarked that the wild cat was now not rare 
in the north of Scotland, and a good account of its increase and 


spread was given by the late Mr. J. G. Millais in 'The Times' of 26 
October 1926. According to that article the wild cat reappeared in 
Inverness-shire, from north of the Caledonian Canal, about 1912 
and, after becoming established round Lochs Ericht and Laggan, 
spread farther south into Perthshire, where one was killed at 
Murthly, twelve miles from Perth, in 1925. 

"Mr. Kinnear further stated that owing to the increase of tree 
planting the safety of the wild cat appeared to be assured, as it had 
taken to the young plantations on account of the rabbits and, where 
the plantations were under the charge of the Forestry Commission, 
the cats were encouraged, as they helped to keep down the rabbits." 
(Kinnear, 1934, p. 68.) 

"The War granted a respite to the Carnivora, and the Wild Cat, 
which but for that event would probably have been exterminated by 
now, increased in numbers. But now the persecution of this animal 
and other carnivores is in full swing again; and apart from a slight 
possibility of help coming from the third cause of change, dealt 
with below [planting of thousands of acres in the Highlands with 
conifers], it is probable that the Wild Cat will be brought to the 
verge of extinction again before long." (Hinton, 1935, pp. 33-34.) 

Peloponnesian Wildcat 


[Felis catus] morea Trouessart, Cat. Mamm., quinq. suppl., fasc. 1, p. 273, 
1904. (Based upon the "Felis catus ferus L. var. e Morea" of Reichenbach, 
Vollstandigste Naturgeschichte, Raubsaugethiere, p. 362, 1852, ex Bory 
de Saint-Vincent, Exped. Sci. Moree, atlas, ser. 3, zool., pi. 1, A, 1833; 
type locality, as restricted by Harper (1940, p. 194), "above Dragomanou, 
near Mt. Diaphorti, west central Morea (Peloponnesus), Greece.") 

FIGS.: Bory de Saint-Vincent, Exped. Sci. Moree, atlas, ser. 3, zool., pi. 1, A, 
1833; Reichenbach, Praktisch-gemeinniitzige Naturgeschichte, Kupfer- 
sammlung, pt. 1, Raubsaugthiere, pi. 80, fig. 639, 1837(?). 

This form of southern Greece, while evidently less common than 
formerly, does not seem to be threatened with extinction and is 
included here chiefly for the purpose of rounding out the picture 
of the European Wildcats. 

It differs from F. s. silvestris in its generally isabelline coloration, 
in the absence of distinct stripes on the sides, and in having the 
black rings on the tail straight and clearly defined; lateral stripes 
replaced by irregular brownish-rufous marblings; feet unspotted 
(Trouessart, 1910, p. 100). 

Bory de Saint-Vincent states (1836, vol. 1, p. 396) that among 
the oak-dotted pastures of the type locality near Mount Diaphorti 
the Wildcats occur in larger numbers than elsewhere. Here he 
collected the type specimen from the high branches of an oak. 


According to Geoffrey (in Bory de Saint-Vincent, 1833, vol. 3, 
pt. 1, zool., p. 13), the animal is very common in certain mountain- 
ous parts of Arcadia, especially in the Canton of Karytaena and on 
the slopes of Mount Diaphorti. It is destructive to poultry, small 
birds and mammals, and partridges. 

The Game Department, Greek Ministry of Agriculture, reports 
(in litt., October, 1936) a general decrease in the numbers of Wild- 
cats in Greece. 

[The Spanish Wildcat (Felis silvestris tartessia *) inhabits the 
Iberian Peninsula south of the Douro and the Ebro, and still 
abounds in the wilder parts. While interbreeding with feral Domestic 
Cats probably takes place, no evidence of it has been found. (Ca- 
brera, 1914, pp. 205-206.) 

The Caucasian Wildcat (Felis silvestris Caucasians 2 ) occurs in 
all the mountain forests and in the greater part of the forested 
lowlands of the Caucasus region. It is generally reported as very 
common. (Ognev, 1930, p. 58.) ] 

Cretan Wildcat. Chat sauvage de Crete (Fr.) 


Felis ocreata agrius Bate, Proc. Zool. Soc. London 1905, pt. 2, p. 317, 1906. 
(Type skin bought in the bazaar at Khania, Crete.) 

This species, which is confined to the island of Crete, may be in 
danger of extinction by "dilution," consisting in this case of inter- 
breeding with feral Domestic Cats. 

The general color is yellowish gray; no black markings on body 
or legs, but indications of brownish shoulder stripes and dorsal 
stripe ; tail with black tip and two or three black subterminal rings ; 
ear blackish at tip (Miller, 1912, p. 470) . 

Raulin (1869, p. 1033) records the species from the woods of the 
lower zones. 

"Hybrids between F. o. agrius and the domestic cat of the island 
appear to be not uncommon, and this can easily be accounted for 
by the fact that formerly small villages were often totally deserted 
for a considerable time, or possibly entirely, during the insurrections 
which occur so frequently in Crete, when the cats, as well as the 
villagers, are forced to take to a life in the hills. Skins of these 
hybrids, which are generally of large size like the true wild race, 
may often be seen hanging up in the bazaars at Khania and Candia." 
(Bate, 1906, p. 318.) 

^ Felis tartessia Miller, Ann. Mag. Nat. Hist., ser. 7, vol. 20, p. 397, 1907. 
("Coto Donana, near Jerez de la Frontera, [Huelva,] Spain.") 

2 Felis catus caucasicus Satunin, Mitteil. Kaukas. Mus., vol. 2, pts. 2-4, pp. 
154 (Russian) and 316 (German), 1906. (Caucasus region.) 


The same author (in Trevor-Batty e, 1913, p. 255) refers to the 
Cretan Wildcat as "not uncommon in the island." 

The Game Division, Forestry Department, Greek Ministry of 
Agriculture, seems to consider (in Hit., March, 1937) that this animal 
is derived from Domestic Cats which the inhabitants left when they 
had to flee during the war of independence, and also from those 
left by the departing Turks when the exchange of populations took 
place after 1922. 

It may be added that Pocock (1907, pp. 151, 160) evidently 
believes that the type specimen was a Domestic Cat or a feral repre- 
sentative of one. 

Cor si can Wildcat. Chat sauvage de Corse (Fr.) 

FELIS BEYI Lavauden 

Felis reyi Lavauden, C. R. Acad. Sci. [Paris], vol. 189, p. 1023, 1929. ("Foret 
d'Aunes des bords de la lagune de Biguglia (Sud de Bastia)," Corsica.) 

This Wildcat is included because of its interest as an insular form 
and because of the generally uncertain future of the Wildcats of 
Europe, rather than on account of any definitely recorded decrease. 

It is smaller than Felis silvestris silvestris; pelage very dark, with 
a darker, rather indistinct dorsal stripe; hind feet with black marks 
like those of African Wildcats; back of the ear dark brown. Head 
and body, 580 mm. ; tail, 270 mm. Weight, 2 kg. (small females) to 
5 kg. (large males). (Lavauden, 1929, pp. 1023-1024.) 

No Wildcat had been recorded from Corsica before 1929, and 
only three specimens have been studied so far, but the species is 
not extremely rare. It is found throughout Corsica in the high 
mountains, the forests, the thickets of the hills, and the shrubbery 
of the plains. The Corsican hunters do not bother to seek the animal 
because of the low value of its fur. (Lavauden, 1929, p. 1024.) 

Sardinian Wildcat. Chat sauvage de Sardaigne (Fr.). Gatto 
selvatico di Sardegna (It.) 


[Felis libyca] var. sarda Lataste, Act. Soc. Linn. Bordeaux, vol. 39, p. 231, 

1885. ("Sarrabus (Sardaigne).") 
FIGS.: Martorelli, 1896, pis. 1, 2. 

While little information is at hand concerning the numerical 
status of this species, it is possibly being subjected, like other 
Wildcats of Europe, to the process of extinction by "dilution" in 
addition to direct persecution. 

It differs from Felis silvestris in its shorter fur and more slender 
tail (hairs at middle averaging about 30 mm. instead of 40 mm.) ; 


hairs of median dorsal line slightly elongated and stiffened; dark 
markings obsolete, the back and sides grayish or brownish, without 
definite stripes; back of ear yellowish clay-color, the tip black; tail 
with well-defined black tip. Head and body, 600 mm. ; tail, 300 mm. 
(Miller, 1912, pp. 468, 470.) 

While this species was originally described from Sardinia, the 
same animal is recorded by Martorelli (1896, p. 266) from the 
Maremma of Tuscany in western Italy. 

It is found throughout Sardinia but is not numerous there, and 
in legislation is rated as harmful (Laboratorio di Zoologia Applicata 
a Caccia, in litt., September, 1936). 

The animal of Tuscany is said to be not rare (Colosi, 1933, p. 56) . 
It is considered a fierce destroyer of hares, pheasants, and other 
game and consequently is much persecuted (Martorelli, 1896, p. 279) . 

European Lynx. Lynx (Fr.). Luchs (Ger.). Lince (Sp., It.) 

LYNX LYNX LYNX (Linnaeus) 

[Felis] Lynx Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 43, 1758. (Near Upsala, 


SYNONYM: Felis borealis Thunberg (1798). 
FIGS.: Wolf, 1867, ser. 2, pi. 6; Elliot, 1883, pi. 39; Martin, 1910, p. 117, fig. 34; 

Colosi, 1933, p. 41. 

The Lynx has suffered rather serious depletion of numbers in 
its European range, and even total extermination in some of the 
countries (Britain, France, Denmark, Switzerland, Italy, Austria, 
Hungary). In northern Asia, however, its status remains much 
more satisfactory, especially in those areas where the human popu- 
lation is still sparse. 

The form is heavier than in Felis silvestris; the legs are relatively 
longer, the feet more robust, and the tail shorter; upper parts and 
sides varying from yellowish brown to brownish gray; back and 
sides never thickly spotted; cheeks not conspicuously whiskered; 
ears conspicuously tufted at tip (Miller, 1912, p. 472). Head and 
body, 800-1,200 mm.; tail, 190-220 mm. (Didier and Rode, 1935, 
p. 289). 

The range of this Lynx includes the forested portions of Europe 
and Asia: north to the tree limit; south to France, northern Italy, 
Yugoslavia, Greece, Bulgaria, the Ukraine, central Russia, the Altai 
Mountains, northern Mongolia, northern Manchuria, and northern 
Korea ; east to Sakhalin and eastern Siberia. 

"The European Lynx, Felis lynx, was ... an inhabitant of 
Britain in the Pleistocene age, and survived until recent times, and 
may even have lingered into the historic period" (Millais, 1904, 
p. 168). 



Although the French authors record the Lynx from the Pyrenees 
(where it is now probably exterminated), Cabrera suggests (1914, 
p. 210) that the animal formerly occurring there was the Spanish 
Lynx (Lynx pardellus) . Trouessart in 1884 (p. 229) considered the 
European Lynx still present in the Alps, the Jura, and the Pyrenees. 
There is a record (the last?) for the Jura in 1834 (Martin, 1910, 

FIG. 29. European Lynx (Lynx lynx lynx) 

p. 118). Didier and Rode (1935, pp. 290-291) cite records from the 
French Alps as late as 1907, 1913, and 1922, but conclude that the 
species has probably disappeared from the entire country. 

In Germany, for several centuries past, the Lynx has occurred 
only as an occasional straggler. From 1773 to 1796 five were shot 
in the Thuringian Forest. A few were taken in Upper Silesia at the 
beginning of the nineteenth century. Two were shot in the Harz 
Mountains in 1817 and 1818. (Blasius, 1857, p. 176.) The species 
was exterminated in Pomerania in 1738; in Westphalia in 1745; in 
Gotha in 1819; in Bavaria in 1850. In East Prussia several were 
taken about 1870, and the animal still occurs frequently on the 
eastern boundary. (Krumbiegel, 1930, p. 6.) One was taken in Thur- 
ingia in 1843, and one in Wiirttemberg in 1846 (Internationale Ge- 
sellschaft zur Erhaltung des Wisents, in Hit., October, 1936). At 


present the law gives it absolute protection (Reichsstelle fiir Natur- 
schutz, in Hit., October, 1936) . 

In Denmark the species is known only from Stone Age and Bronze 
Age remains (Winge, 1908, p. 117) . 

In Switzerland it abounded during the seventeenth century, and 
numerous captures were made up to the early part of the nineteenth 
century. Thereafter it suffered a pronounced decrease. In the 
1860's it was still found, but only occasionally, in Grisons, Ticino, 
and Valais (one record in 1867). (Fatio, 1869, p. 280.) One was 
killed in the Engadine in 1872. The species is now extinct in Switzer- 
land. (Federal Forest, Game, and Fish Inspection, Berne, in litt., 
March, 1937.) 

The species may be considered extinct in Italy, the last specimens 
having been killed in Piedmont in the second half of the last century 
(Laboratorio di Zoologia Applicata a Caccia, in litt., September, 
1936). De Beaux (1932, p. 9) speaks of the forest of Langhe in the 
Maritime Alps, in the province of Cuneo, as its last refuge. 

In Austria the Lynx is entirely exterminated. In Carinthia one 
was killed in 1848, and another was seen in 1878. In Lower Austria 
it was fairly distributed up to the middle of the last century. In the 
Tyrol it was very common in the sixteenth century, but during 
the next century it decreased decidedly, and at the beginning of the 
nineteenth century it was said to be not very rare ; the last one was 
killed at Graun in 1873. In Vorarlberg the last one was killed in 
1918. (G. Schlesinger, in litt., March, 1937.) 

Before World War I the Lynx was found everywhere in the Car- 
pathian forests of Hungary (now Czechoslovakia) (J. Schenk, in 
litt., November, 1936) . In the higher elevations of the Tatra Range 
(Czechoslovakia and Poland) there are a few Lynxes (Maurice, 
1927, p. 21). (See also under Poland.) 

In the present Hungary the species is not found (J. Schenk, in litt., 
November, 1936) . 

During the past century the Lynx was exterminated throughout 
Yugoslavia except in the high mountain region of Shar Planina, in 
southern Serbia, where four or five specimens are killed every year 
(M. Hirtz, in litt., December, 1936). 

In Greece it occurs in Epirus, Thessaly, Macedonia, and Thrace. 
No decrease has been observed. (Game Department, Greek Ministry 
of Agriculture, in litt., October, 1936.) 

In Bulgaria "the lynx is about gone, though I saw two fine skins 
last week at a fur shop in Varna, on the Black Sea, and one in the 
peasant market in Sofia last year." A reserve for the Lynx in the 
Pirene Mountains is advocated. (H. W. Shoemaker, in litt., June 
30, 1932.) According to the Bulgarian Game Association "Sokol" 
(in litt., February, 1937) , the last specimen was killed in 1907. The 


cause of its disappearance was the diminution of the forests (espe- 
cially the virgin forests) . 

In Rumania the Lynx is an increasingly rarer species. Its dis- 
tributional area is the coniferous-forest zone narrower on the 
western slope of the eastern Carpathians (Nasaud district), and 
broader on the eastern slope (Bukowina) . It is more common in the 
southern Carpathians (Bunzenlander Mountains, Fagaras Moun- 
tains, etc.). (Calinescu, 1930, p. 366.) 

In ancient times, when Poland was covered with large forests, the 
Lynx was found everywhere. It still exists in the Carpathians, espe- 
cially in the eastern Polish part, and also in the great forests of 
northeastern and eastern Poland. The estimated number is about 
400. During the hunting season about 25 specimens are killed 
annually. Hunting is forbidden from March 1 to December 31. (M. 
Siedlecki, 1 in'litt., October, 1936.) In the proposed International 
Tatra Park (Poland and Czechoslovakia) the Lynx will be kept 
under absolute protection. The chief aim of another proposed Inter- 
national Park (Poland, Czechoslovakia, and Rumania) is the crea- 
tion of a breeding ground for the Bear, Lynx, Wildcat, Wolf, Stag, 
etc. (Benedyct Fulinski, MS., 1933.) 

In Lithuania the Lynx is almost exterminated. Since the Great 
War about ten specimens have been killed. Hunting is forbidden. 
(T. Ivanauskas, in litt, November, 1936.) 

In Latvia the species has decreased considerably but is still found 
in the large forests. The following numbers have been reported 
by the Forest Department: 74 in 1925; 49 in 1932; 59 in 1935; 78 
in 1936. (N. von Transehe, in litt., February, 1937.) The animals 
are found especially in the northeastern part of the country, where 
hunting is restricted; in other parts no protective measures have 
been adopted. Twenty to thirty years ago the stock was about 
300-400, but for economic reasons, and in order to protect useful 
game, the numbers have been reduced intentionally, and they are 
now confined to a certain part of the forests, where they are pro- 
tected. (Forest Department, Latvia, in litt., March, 1937.) 

In Estonia the Lynx is only a visitor, coming in from Russia, and 
is given no protection (Zoological Institute, University of Tartu, 
in litt., October, 1936) . 

i "Michel Siedlecki, Professor of Zoology in the University of Cracow, was 
... an enlightened apostle of Fauna Preservation. . . . 

"Michel Siedlecki was done to death in one of Germany's most notorious 
concentration camps. . . . 

"In . . . international relations Siedlecki inspired among his colleagues both 
respect and affection. One wonders with what feelings the German colleagues 
with whom he collaborated in the International Council for the Exploration 
of the Sea, as well as in the International Committee for Bird Preservation, 
reflect upon the death, brought about through the calculated brutality of their 
Government, of a loyal colleague who was so recently their guest." (Jour. Soc. 
Preservation Fauna Empire, n. s., pt. 39, pp. 15-16, 1940.) 


The species was formerly common in the Norwegian forests 
north to about latitude 65 30'. A quarter of a century ago Collett 
remarked on a great decrease. Since it preys upon the more im- 
portant smaller game and on small cattle, it is hunted throughout 
the country and is not protected by law. (Hj. Broch, in Hit., Decem- 
ber, 1936.) Up to about 1875 it was generally distributed in the 
woodlands of Norway. A small stock remains in some places in 
Fosen, Namdalen, and South Helgeland. In southern Norway it is 
doubtful if there is any resident stock, but now and then some 
stragglers may appear. Up to the present, bounties have been paid 
on the Lynx, and it is doubtful if its total extermination in Norway 
can be prevented. (Director of Forestry, Norwegian Ministry of 
Agriculture, in litt., January, 1937.) 

About a century ago the Lynx was rather common throughout 
Sweden except in the northernmost provinces and in some of the 
southernmost. From 1827 to 1839, 3,224 Lynxes were killed an 
annual average of 248; in 1844, 250; and in 1845, 273. The annual 
average was about 175 in 1856-60; 121 in 1865-69; 105 in 1871-75; 
67 in 1876-80; 24 in 1881-85; 35 in 1891-1900; 10 in 1901-05; 17 
(all in the four northernmost provinces) in 1906-10; 11 in 1921-25. 
In the southern provinces the species was practically exterminated 
before 1870; in the middle provinces it was found in diminished 
numbers in the 1890's. The increase in numbers killed in the 1890's 
and in 1906-10 was due to livelier persecution in the more northern 
provinces. 1 In 1926 and 1927, 7 Lynxes were killed. Since then the 
species has been protected by law to the extent that it may not be 
killed on public domains, and, if killed on private lands, it is never- 
theless crown property, so that the profit motive is eliminated. The 
crown domains are very extensive, especially in the northern prov- 
inces, and there are some state forests, especially in Westerbotten, 
where the animals seem to thrive and to increase somewhat. There 
are a few in Angermanland and perhaps also in Jemtland, while 
stragglers have been traced in some other provinces. (Einar Lonn- 
berg, in litt., October, 1936.) 

The following information on the Lynx's range and status in 
Russia and Siberia is derived from Ognev (1935, pp. 206-214) : 
The range extends entirely across Russia from east to west; in the 
north it reaches the tree limit on the Kola Peninsula and at other 
points near the Arctic coast; in the south it reaches Podolia, the 
southern part of the Western and the Moscow Areas, the district 
of Penza, and the former Governments of Kazan and Orenburg. 
Within this range its numbers vary considerably but are evidently 

1 Possibly these increases represented peaks in a periodic fluctuation, such 
as is evident in the case of the Canada Lynx; 1895-97 and 1905-06 were peak 
periods for the latter species (cf. Seton, 1929, vol. 4, p. 711). Ed. 


much less than formerly. In White Russia 76 animals were taken 
in 1924-25, but only 16 in 1925-26. In the Western Area 156 were 
killed in 1928-29, 43 in 1929-30, and 47 in 1930-31. According to 
Milovanowicz (1925), 50 are taken annually in the Petchora region. 
In the Vichegda Basin 3 were taken in 1929, 2 in 1930, and 27 
in 1931. 

In Siberia the Lynx ranges north approximately to the tree limit 
and south to northern Russian Turkestan (Irtish River and Zaisan 
Nor) and to the Mongolian and Manchurian boundaries, apparently 
avoiding the Arctic tundra on the one hand and the steppes of Rus- 
sian Turkestan on the other. (In far eastern Siberia, beyond the 
Verkhoyansk Mountains, the typical subspecies seems to be replaced 
by Lynx lynx wrangeli Ognev.) 

According to W. G. Heptner (in litt., December, 1936) , the Lynx 
is quite common in Siberia but rarer in Russia. Hunting is allowed 
the whole year, but in certain parts of Russia and in western Siberia, 
where the numbers of the animal are small, hunting is limited to 
certain open seasons. 

The species occurs in the mountains of northern Mongolia, from 
the Altai Range eastward (Ognev, 1935, p. 214) . A specimen from 
15 miles northeast of Urga, Mongolia, is recorded by G. M. Allen 
(1929, p. 14) under the name of L. I. isabellina. The species also 
occurs in the forests of Manchuria; it is rare in northern Kirin 
(Sowerby, 1923, p. 37) and in the southern part of the Little Khin- 
gan Mountains (Ognev, 1935, p. 213). It is generally considered 
rare in Sakhalin (Kuroda, 1928, p. 226; Miyoshi, in Skottsberg, 
1934, p. 411). 

Economics. The Lynx attacks game as large as the Red Deer 
and the Roe Deer, and it has been known to slaughter 30 sheep in a 
single night. Generally it feeds on the smaller game hares, mar- 
mots, small rodents, and birds of all kinds. On occasion it does not 
fear to attack man himself. (Trouessart, 1884, p. 230.) 

In Sweden, "when the lynxes were numerous, they sometimes killed 
sheep and perhaps also reindeer. They are therefore like all other 
carnivorous animals especially hated by the Lapps. The lynxes 
were also destructive to the hares, when they were common. The 
reason why they were so much hunted was, however, chiefly because 
it was regarded as a good sport, and the value of the skin was also 
attractive." The species is now protected in Sweden, and the govern- 
ment pays for the damages committed, "if they are not due to 
carelessness of the owners of the domestic animals killed." (Einar 
Lonnberg, in litt., October, 1936, and January 18, 1933.) 

"The Norwegian peasants believe that if a person wears a neck- 
lace made of the fore-claws of a lynx it will preserve him from 
spasms and the cramp. It would appear that the Russians entertain 


a somewhat similar superstition, for when they sell lynx skins to the 
Chinese, they charge a much higher price for them if the fore-claws 
are included." (Bowden, 1869, p. 14.) 

In the Polish forests "the lynx and the wolf alone are capable 
of killing the elk" (Korsak, 1934, pp. 78-79). 

"The Russian naturalists Von Schrenk and Radde inform us that 
the natives of Amoorland esteem the flesh of this animal as a great 
delicacy, and that the furs which are obtained by the hunters in 
this part of Asia mostly pass into Chinese hands, being much trea- 
sured by the high officials of the Celestial Empire" (Sclater, in Wolf, 
1867, text to pi. 6). 

Tibetan Lynx 


Felis isabellina Blyth, Jour. Asiatic Soc. Bengal, vol. 16, p. 1178, 1847. ("Tibet.") 
FIGS.: China Jour., vol. 23, no. 3, pi. facing p. 172, 1935, and vol. 25, no. 5, 
pi. facing p. 288, 1936; Schafer, 1937, pi. facing p. 177. 

The Tibetan Lynx seems to be a moderately rare animal, and 
while it is protected to some extent by the remoteness of its haunts, 
its fur is in considerable demand. 

It differs from the European Lynx in its pale sandy gray or 
isabelline coloring, and in the relative shortness of the hair on the 
toes. Head and body, 837 mm. ; tail, 196 mm. ; weight, about 60 Ib. 
(Blanford, 1888-91, p. 90.) 

"This race inhabits the plateau of Eastern and Western Tibet, 
and certainly extends into Baltistan; but its exact geographical 
limits are impossible to define .... 

"Throughout its habitat, so far as accounts go, the Tibetan lynx 
is a rare animal, seldom seen, and still more rarely shot. . . . The 
Tibetan hares and blue pigeons form the chief prey of the lynx in 
Ladak, although it also levies toll on the smaller domesticated ani- 
mals of the Tatars." (Lydekker, 1900, pp. 326-327.) 

"This animal is rarely encountered and consequently the exact 
limits of its habitat are somewhat conjectural, but I fancy that it is 
almost identical with that of Ammon. . . . They are savage ani- 
mals and do not hesitate to attack sheep and goats, sometimes 
working considerable havoc." (Burrard, 1925?, p. 241.) 

"Lynx skins . . . are brought in from the Thibetan regions to 
the north and west, to Sungpan [Szechwan] , where they find a ready 
market among the wealthy Chinese. . . . They sell in Sungpan 
for 5 to 7 taels each." (Wilson, 1913, vol. 2, p. 181.) 

"The Isabelline Lynx ... is fairly common in the mountainous 
regions along the Chinese-Tibetan border. . . . The lynx supplies 
the fur market of this country [China] with one of its best furs, the 


coat of this animal being long, thick and soft." (Sowerby, 1936, 
pi. facing p. 288.) 

Ognev (1935, pp. 215, 231-232) reports it from Kansu, Koko Nor, 
the Zaidam region, the Nan-Shan, Altyn-Tagh, Kwen-Lun, Tian- 
Shan, Borokhoro, and Bogdo-Ola ranges, Kashgar, the Tarim Basin, 
and Lob Nor. 

The systematic status of the Lynxes inhabiting various mountain- 
ous areas from northwestern India and southern Russian Turkestan 
to northern Persia seems to be in doubt; I am unable to determine 
what subspecific name or names should be applied to them. This 
vast region lies directly between the ranges of the Tibetan Lynx 
and the Caucasian Lynx. 

The animal of Gilgit, in the valley of the Indus, has a more 
rufous coloring than that of Tibet (Lydekker, 1900, p. 326) . 

According to Ognev (1935, pp. 214-215), the Lynx is found in 
various localities of Russian Turkestan, including the Pamirs, the 
western Tian-Shan, Semiretchie, the Chu River, the Kara Tau, the 
Talassk Alatau, the Samarkand region, and the Kopet-Dagh (where 
it is rare). It occurs in northern Afghanistan and doubtless in the 
Persian provinces of Gilian, Mazanderan, and Astrabad; possibly 
also in the mountains of Khorassan (lat. 37 N.). 

W. G. Heptner states (in litt., December, 1936) that the Lynx 
occurs in small numbers in the mountains of Turkestan. Hunting 
is limited to certain seasons in the mountains of Uzbekistan, and is 
forbidden on the Kopet-Dagh. 

Barbary Lynx. Lynx caracal (Fr.) 


Felis Caracal . . . Var. algira Wagner, Reisen Regentschaft Algier, vol. 3, 
p. 76, atlas, pi. 4, 1841. (Vicinity of Algiers (op. tit., p. 62).) 

FIGS.: Buffon, Hist. Nat., vol. 9, pi. 24, 1761; M. Wagner, 1841, atlas, pi. 4; 
Loche, 1867, pi. 2. 

This Caracal is evidently becoming increasingly scarce as the 
years roll by. 

General color nearly uniform, between cinnamon-orange and 
reddish cinnamon; paler about the eyes, on the lips, and on the 
lower parts; a blackish spot on each side of the mouth; ears ex- 
ternally black, sometimes mixed with white hairs, terminal tuft 
black. Head and body, 717 mm.; tail, 284 mm. (Cabrera, 1932, 
pp. 171-172.) 

The species as a whole is "widely distributed in suitable localities 
from South Africa to Egypt and Morocco, and from Palestine to 
India" (Flower, 1929, p. 83). Only the North African subspecies 


(algirus) calls for attention here. It ranges from the Gulf of Gabes 
to the Atlantic coast, and south to Senegal. In Algeria and Tunisia 
it reaches the Mediterranean coast. Apparently it is not represented 
in Tripolitania. (Cabrera, 1932, pp. 172-173.) 

Tunisia. It is rare in the north but a little more common in the 
center and south (region of Feriana, Djebel-Selloum, Djebel-Bou- 
Hedma). It can be shot rather easily in its usual habitat among 
alfa grass, and it could be successfully chased with hounds, for its 
gait is not very rapid. (Lavauden, 1932, p. 7.) 

FIG. 30. Barbary Lynx (Caracal caracal algirus). From specimen in 
Philadelphia Zoo. 

Algeria. The Barbary Lynx does not occur commonly in the 
vicinity of Algiers, whence occasionally specimens are brought to 
market (M. Wagner, 1841, vol. 3, p. 62). 

Loche (1867, p. 41) reports for the period 1840-42 that it ranges 
throughout Algeria, where it is rather numerous. Specimens are 
recorded from Birkadem, Arba, and Djelfa. It is also met with near 
Coleah, Lac Halloula, and elsewhere. 

Known in Barbary from the first explorations, it has been recorded 
by Shaw, Poiret, and others. Two specimens were taken in the 
vicinity of Laghouat, and one of the animals was seen between 
Haidra and Tebessa, Algeria. Without being very abundant, the 
species seems rather widespread, and skins are frequently seen in 
the saddlers' shops of Algiers and Constantine. (Lataste, 1885, 
p. 225.) 


Heim de Balsac (1936, p. 179) records it from the valley of the 
Saura, near Beni-Abbes. 

Morocco. "The Lynx ... is found in wooded districts, and is 
sometimes brought alive to Mogador" (Leared, 1876, p. 304). It 
seems to be lacking in Yebala, but it undoubtedly exists more to 
the south, and it is reported as still living in the interior of the 
Rif (Cabrera, 1932, pp. 172-173) . In the Zaian district it is found 
from time to time, and its skin is frequently seen in the market. One 
was living in captivity as late as 1931. (Carpentier, 1932, p. 17.) 
Specimens are recorded from Gara de Debdou, Matarka, Oued 
Charef , region of Berguent (Laurent, 1935, p. 349) . 

Gambia. "This splendid animal is to be seen some 150 miles up 
the [Gambia] river; being swift and cunning, very few are trapped 
or shot" (E. Johnson, 1937, p. 63). The local form does not seem 
to have been subspecifically determined. 

The Cheetahs (genus Acinonyx) 

These animals are also known as Hunting Leopards. Additional 
names in various Continental languages are: Guepard (Fr.), Gep- 
pard (Ger.), Ghepardo (It.), and Onza (Span.). 

The common species of Cheetah (Acinonyx jubatus) , which has 
been divided by various authors into approximately a dozen differ- 
ent forms, has become rather rare over a large part of its enormous 
range in Africa and Asia, while remaining moderately common in 
some areas. Another species, the King Cheetah (A. rex), has been 
described from a restricted area in Southern Rhodesia. For the 
sake of completeness, all forms that are more or less recognizable 
will be treated in the following accounts. In scarcely any case can 
the distributional limits of the subspecies be stated precisely, owing 
to the incompleteness of our present knowledge. 

The Cheetahs are distinguished from all other members of the 
cat family (Felidae) by the absence of claw-sheaths (Pocock, 1916, 
p. 426). In size and form they suggest a long-legged and slender- 
bodied Leopard; but their markings are solid spots instead of ro- 
settes, as in the Leopard. The hair of the neck is elongated to form 
a slight mane. (Lydekker, 1900, p. 328.) 

The geographical distribution of the Cheetahs as a genus is very 
similar to that of the Lion. In Africa it includes chiefly the arid 
or semiarid areas of the South, East, and North in fact, most of 
the continent outside of the rain forests of the West African sub- 
region and the humid, forested areas of southeastern Africa. In 
Asia, likewise, it includes more or less arid areas from India and 
Russian Turkestan to Syria, Palestine, and Arabia. 


Economics and conservation. "The Cheetah has been for cen- 
turies the playing thing of princes, Asiatic, African and European. 
When taken young it is easily tamed and trained to show its won- 
derful speed. A couple of cheetahs were sent as a present to Ana- 
stasius, Emperor of the East, at Constantinople in A. D. 439 from 
whence the sport reached Italy and obtained an enormous vogue 
among the wealthy, extravagant grandees of the great City States. 
The Turks, Moors and Persians alike used cheetahs in hunting. In 
India the Cheetah is usually taken blind-folded in a cart to the 
scene of the hunt. In the proximity of a herd of antelope it is un- 
hooded and slipped from its leash: a short crouching stalk and a 
few bounds of great length and rapidity and the hunt is over 
the quarry has escaped or the Cheetah holds it in a strangle-hold 
by the throat, till the keeper comes up and having cut the captive's 
throat rewards the captor with a drink of warm blood collected in 
its accustomed feeding bowl." (Anonymous, 1935, pp. 148-149.) 

(For an interesting account of hunting with Cheetahs in the 
Middle Ages, see Yule's Marco Polo, ed. 3, vol. 1, pp. 397-398, 

"So far as I have heard, . . . this animal has not been known to 
breed in captivity" (Blanford, 1888, p. 93) . 

Probably the disappearance of the Cheetah in the Cape Province 
and its decrease in North Africa and in Asia are closely linked with 
the general decline in those regions of the various antelopes which 
constitute its principal prey. It also attacks calves, sheep, and goats 
to some extent, and thereby comes into conflict with man. However, 
"there are very few recorded cases of cheetah attacking human be- 
ings or taking the offensive, even when wounded or in defence of 
their cubs" (Shortridge, 1934, p. 107) . Like all of the larger carni- 
vores, it is evidently shot "on general principles." According to 
Shortridge (1934, p. 107), it retreats more rapidly than the Leopard 
before European settlement. 

In Northern Rhodesia "Cheetah and other animals take their toll 
of the vast numbers of Black Lechwe, but these natural enemies do 
more good than harm" (David Ross, in litt., May 3, 1933) . 

Fortunately the Cheetah is reported present in a considerable 
number of the African game reserves: White Nile Reserve and one 
near the Ethiopian border in the Sudan; reserve between the Gash 
and Setit Rivers in Eritrea; Northern and Southern Reserves in 
Kenya; Katavi, Mtandu, Lake Natron, Ngorongoro, Northern Rail- 
way, Saba, Selous, and Serengeti Reserves in Tanganyika; Matupo 
Reserve in Mozambique; Kruger National Park in Transvaal; 
Bechuanaland Reserve in Bechuanaland Protectorate; and Namu- 
toni Reserve in South- West Africa. Such reserves doubtless afford 


the species its best chance of survival in countries that continue to 
be opened up to settlement and cultivation. 
Cheetahs make charming pets, and may be led about on a leash. 

South African Cheetah 


Felis jubata Schreber, Saugethiere, vol. 3, pi. 105, 1776; p. 392, 1777. (Cape 

of Good Hope, South Africa.) 
SYNONYMS: Felis lanea P. L. Sclater (1877); lAcinonyx guttatus obergi 

Hilzheimer (1913). 
FIGS.: Proc. Zool. Soc. London 1877, pi. 35 (incipient albinism); Hilzheimer, 

1913, p. 289, fig. 2 (A. j. obergi) 1 

The South African Cheetah has virtually disappeared from the 
Cape Province. Elsewhere in its huge range it has managed to sur- 
vive in varying numbers, but in general it is obliged to retreat before 

The general color is ochraceous-yellow, covered with round black 
spots; below almost white, with longer hair and indistinct spots; a 
black line from the anterior angle of the eye to the edge of the lip ; 
ear with a black patch posteriorly and a tawny tip; chin white, 
unspotted; chest whitish, spotted; spots on the tail tending to form 
6-8 imperfect rings toward the tip. (W. L. Sclater, 1900, p. 47.) 
Total length, about 6 feet 8 inches; tail, about 2 feet 6 inches; 
shoulder height, about 2 feet 8 inches. 

All Cheetahs (except A. rex Pocock) occurring from the Cape 
Province north to Angola, Belgian Congo, Northern Rhodesia, and 
Nyasaland will be included provisionally under the present sub- 
species. This range corresponds roughly to the Southwest Arid 
District, the Southeast Veld District, and the Rhodesian Highland 
District of Chapin (1932, p. 90), or to the Southeast Veldt District, 
the Kalahari and the Damara Arid Districts, and the Rhodesian Sa- 
vanna District of Bowen (1933, pp. 256, 259-260) . 

"In the [Cape] Colony it is found sparingly in the western and 
midland districts : north of the Orange River it is common in German 
territory [South-West Africa], the Kalahari and Bechuanaland, and 
exists in Rhodesia, the Transvaal, Zululand and Natal, though now 
very rare in the latter Colony, and found only in the Drakensberg 
range" (W. L. Sclater, 1900, p. 48) . 

In South-West Africa the Cheetah is widely distributed. It is 
considered quite plentiful in the eastern sand-veld region, scarce 
in the Kaokoveld, sparsely distributed in the Caprivi, and not un- 
common in the Namutoni Game Reserve. It is recorded from various 

i The following additional figures of Cheetahs are of undetermined subspecific 
identity: Wolf, 1861, pi. 13; Lavauden, 1924, p. 10, fig.; Leplae, 1925, p. 46, 
fig.; Malbrant, 1936, pi. 1, upper fig. 


parts of Omaheke. It is apparently scarce in the vicinity of the 
Orange River, the western and southwestern parts of Great Nama- 
qualand, and the highlands of western Damaraland. Its numbers 
increase in the sand-plain country adjoining Bechuanaland. Con- 
siderable numbers of skins are brought into Windhoek and Keet- 
manshoep annually by natives from this region and from Bechuana- 
land. The species also occurs in small numbers in southern and 
southeastern Angola. (Shortridge, 1934, p. 105.) This author adds 
(pp. 105-107) : 

The Cheetah has almost if not completely disappeared from the Cape 
Province, the Orange Free State, Natal, and the Southern Transvaal, but 
may still be met with in some of the more sparsely populated districts of the 
Northern Transvaal, Zululand, Swaziland, and probably the inland portions 
of Portuguese East Africa. It is widely distributed in Bechuanaland and 
still comparatively plentiful in the central and northern portions of that 
territory. . . . 

The Cheetah is retreating rapidly before settlement in Southern Africa, and 
it is doubtful if there are any to be found to-day south of the Vaal River. . . . 

Cheetah prey mostly upon medium-sized antelope, from steinbok and duiker 
up to the size of impala, springbok, reedbuck, and even cow kudu. . . . 

When opportunity offers they kill sheep, goats, and ostriches, which last 
are driven into wire fences and cornered. 

In the Transvaal the species is considered inimical to man, and 
not a game animal. Thus it is not given any protection. (Austin 
Roberts, in Hit., November, 1936.) 

In the Kruger National Park, Transvaal, "the status of the species 
remains fairly constant; they were never very numerous, and no 
noticeable increase or decrease is reported" (Game Warden's Annual 
Report, 1925?). 

In Southern Rhodesia Cheetahs are sparingly distributed over 
the greater part of the country, but soon retire from inhabited 
areas. They seldom raid domestic stock and consequently do not 
often fall a victim to traps and poison. There is no legal protection, 
but in the recognized game reserves and also in the forest reserves all 
animals are rigidly protected. (Game Warden, Wankie Game Re- 
serve, in Hit., March, 1937.) 

In Northern Rhodesia this "widely distributed species ... is 
absent from the regions of interminable woodland. Usually occurs 
sparingly, though inclined to be locally plentiful." It "is most nu- 
merous in the neighbourhood of open expanses such as the Kafue 
flats, the Batonga and Batoka plateaux, the neighbourhood of Bang- 
weulu, the Chambeshi flats and other similar localities. It appears 
to be absent from a great part of the Luangwa Valley. Family 
parties up to five are frequently reported, and as many as seven have 
been seen together. The cheetah is a very disturbing factor in locali- 
ties where it occurs side-by-side with domestic stock, and is apt to be 


particularly destructive where sheep and calves are concerned." 
(Pitman, 1934, p. 12.) Skins have been obtained in the Mumbwa, 
Namwala, Broken Hill, Mpika, Chinsali, and Petauke Districts 
(op. cit., p. 159). 

In Nyasaland "the Cheetah has so far only been found in the 
Central Province of Angoniland. Even there it is seldom seen, but 
may be more numerous than is believed at present, as it is nearly 
always confused with the Leopard by the natives." (Wood, in May- 
don, 1932, p. 316.) 

In Portuguese East Africa "Cheetahs ... are not very numerous 
in the Zambezi valley, occurring perhaps most plentifully between 
Muterara and the Lupata Gorge, where reed buck and other small 
antelopes are common. I have also seen them in the Mlanje district 
. . . , in the Barue to the south of Tete, and in the open country 
south of the Shupanga Forest." (Maugham, 1914, p. 195.) 

In the Belgian Congo, 20 years ago, the species was comparatively 
abundant over the southern part of the colony, from Kwango to 
Tanganyika. It probably ranged to the northern extremity of Lake 
Tanganyika, wherever grassy stretches, inhabited by small rumi- 
nants, assured it of favorable conditions for existence. The range 
seems to have remained practically the same, except in southern 
Katanga and Lomami, where stock-raising has led to the extermina- 
tion of Cheetahs. It is well to note that these stock farms have 
been established where Cheetahs were particularly numerous. Fur- 
thermore, these animals suffer greatly from hunting with encircling 
fires, and the natives persecute them everywhere to satisfy the 
demand for skins on the part of the European population. The 
Cheetah should be put on the protected list. That would suffice, if 
the customary destructive hunting by the natives could be effectively 
stopped, to assure the recuperation of the species outside the zones 
of stock-raising. (A. J. Jobaert, in litt., November 10, 1936.) 

East African Cheetah 


A[cinonyx\ g[uttatus] ngorongorensis Hilzheimer, Sitz.-ber. Ges. naturf. 
Freunde Berlin 1913, no. 5, p. 290, figs. 3-4, 1913. (Based upon a living 
specimen in the Leipzig Zoological Garden, said to have come from 
"Ngorongoro," south of Lake Natron, Tanganyika Territory.) 

FIGS.: Hilzheimer, 1913, pp. 290-291, figs. 3-4; Roosevelt and Heller, 1914, 
pi. facing p. 244 (raineyi) ; Hollister, 1918, pi. 5, lower fig. (raineyi) ; 
Zammarano, 1930, p. 152, fig. (ngorongorensis?). 

Two other forms were subsequently described from East Africa: 
A. j. velox Heller, Smithsonian Misc. Coll., vol. 61, no. 19, p. 7, 1913 
("Loita Plains, British East Africa"), and A. j. raineyi Heller, 
Smithsonian Misc. Coll., vol. 61, no. 19, p. 9, 1913 ("Ulu, Kapiti 


Plains, British East Africa"). Hollister (1918, p. 151) seems to be 
somewhat doubtful as to the distinctness of these three East African 
forms from each other, and De Beaux (1927, pp. 3-4) does not 
recognize velox or raineyi. The Cheetahs of East Africa, from 
Tanganyika north to Ethiopia and Eritrea, will be treated here 
as a unit. This range corresponds roughly to the Somali Arid District 
and the East African Highland District of Chapin (1932, p. 90), or 
to the Somali Arid District, the North Kenya Savanna District, 
and the East African Highland District of Bowen (1933, pp. 256, 

The ground color of the living type specimen of ngorongorensis 
was isabella yellow-brown; under parts very light isabella, entirely 
without white; spotting of the lower neck very pronounced; cheeks 
grayish, heavily spotted; back of the ear yellow, with a slender 
black basal stripe; chin and lips white; chest and belly unspotted; 
outer side of the limbs heavily spotted as far as the toes; tail with 
three complete rings, tip yellowish white (Hilzheimer, 1913, p. 289) . 
The coloration of this captive specimen may not have been typical. 

The following might serve as a composite characterization of velox 
and raineyi: ground color ochraceous to light pinkish buff; spots 
large, blackish; snout ochraceous to ochraceous-buff ; a black tear 
stripe from eye to mouth; back of ears black basally, tip and inner 
surface buff or pinkish buff; chin and upper throat white to cream- 
buff ; belly cream to cream-buff, with elongate spots ; hind feet more 
or less distinctly spotted; terminal part of tail ringed with black, 
tip whitish (Heller, 1913, pp. 8-10). Head and body, 1,120-1,300 
mm.; tail, 720-800 mm. '(Hollister, 1918, p. 154). 

In East Africa and in South-West Africa the Cheetah seems to 
have survived in more satisfactory numbers than elsewhere in its 
wide range. 

It occurs throughout Tanganyika Territory in varying numbers. 
There is no danger of extinction at present. In five provinces out of 
eight, only two Cheetahs may be killed on a Full Licence, and only 
one on a Minor Licence. (Game Preservation Department, Tangan- 
yika Territory, in litt., December, 1936.) It is "quite numerous in 
Masailand" (Browne, in Maydon, 1932, p. 312). 

In Kenya the Cheetah is fairly common and generally distributed, 
specimens being recorded from the Loita, Kapiti, and Athi Plains, 
Laikipia and Uasin Gishu Plateaus, the flanks of Kilimanjaro, and 
Upper Tana River (Roosevelt and Heller, 1914, pp. 244-249) . There 
are fair numbers in the Southern and Northern Game Reserves (Per- 
cival, 1923, pp. 69-71). There is a decrease in the Native Reserves 
and in the European settled areas; otherwise it is fairly common, 
and it is protected (Game Warden, Kenya, in litt, November, 1936) . 


The Cheetah is sparingly distributed throughout northern and 
eastern Uganda. Not long ago it had a much more extensive distri- 
bution, occurring throughout the savanna regions. Its disappearance 
from many localities is due to the extension of settlement and culti- 
vation. The Cheetah is of sentimental importance and also of 
considerable economic value, being sought after for hunting ante- 
lopes in India. Specimens trained by their parents in the field have 
a local value of 20 to 30. Only one specimen is allowed on a 
Full Game Licence. (Game Warden, Uganda, in litt., December, 

De Beaux (1927, p. 4) records the species from Italian Somaliland, 
Ethiopia, and Eritrea. In the southern plain of Eritrea, especially 
between Barca and Gash, the animal is rather frequent (Zamma- 
rano, 1930, p. 61). 

In British Somaliland "the cheetah is commonest in the thick 
bush country on the edge of the Haud, although it is to be found 
both on Guban and Ogo-Guban" (Drake-Brockman, 1910, p. 22). 

Sudan Cheetah 


Cynailurus Soemmeringii "Riippell" Fitzinger, Sitz.-ber. math.-nat. Cl. Akad. 
Wiss. [Wien], vol. 17, Heft 2, p. 245, 1855. (Based upon a living specimen 
from the Kababish Steppes in the south of the Bayuda Desert, Kordofan.) 

Roosevelt and Heller (1914, p. 249) give the range as "lowlands 
of the Nile Valley, from the Albert Nyanza northward to Kordofan 
and westward to Lake Chad and northern Nigeria." No information 
is at hand as to the exact northern or western limits of this sub- 
species, where it should presumably intergrade with A. j. hecki. 
Its range lies in the eastern portions of the Sudanese Arid District 
and the Sudanese Savanna District of Chapin (1932, p. 90) and of 
Bowen (1933, pp. 256, 258). 

The ground color above is ochraceous or pinkish buff; spots not 
exceeding half an inch in diameter and widely separated ; hind feet 
unspotted. However, according to Malbrant (1936, p. 137 and pi. 1, 
upper fig.), the hind legs are nearly always spotted in Cheetahs of 
the Chad region. 

Roosevelt and Heller (1914, p. 250) record specimens from El 
Dueim on the White Nile and from Lake Chad. "It is a rare animal 
in the Nile district and is seldom secured by sportsmen. . . . Heller 
saw a pair near Gondokoro." 

"Cheetah, although by no means common in the Sudan, are widely 
distributed throughout the country. They are even reported to 
exist as far north as Jebel Tegaru in the north-west corner of the 
Province of Kordofan." (Brocklehurst, 1931, p. 32.) 


Butler (in Maydon, 1932, p. 139) refers to this Cheetah in the 
Blue Nile district as "everywhere a much scarcer beast than the 
Leopard, and rarer in the Eastern Sudan than it is in Kordofan. 
Indeed, on this side of the country I only met it twice, both times 
on the Setit." 

"Heuglin gives the following locality for the Chitah: Southern 
Takah and Eastern Sudan not north of 19 N. ... Ruppell . . . 
mentions Felis guttata as one of the animals hunted ... in the 
western deserts of the Dongola district." (Anderson and de Winton, 
1902, p. 185.) 

In the Ubangi-Shari district of French Equatorial Africa a few 
Cheetahs may occur in the extreme north (Birao), but it is not 
certain. They are not threatened for the moment." (L. Blancou, 
in litt., December, 1936.) 

In the French Cameroons the species is found in the thorn-bush 
country, but is very rare. It does not have any special legal pro- 
tection. (Ministry of Colonies, Paris, in litt., November 7, 1936.) 

In writing of the Chad region, Malbrant says (1936, pp. 137-138) 
that the Cheetah is found in the whole of the Sahelian region of 
central Africa and in the somewhat forested steppes, its southern 
limit being at about lat. 10 N. It lives sometimes solitarily, but 
more often in bands of two to four individuals. The natives of Chad 
do not utilize it for the chase. 

It is found in French Sudan, the Niger Territory, Borku and 
Ennedi, and the desert part of the Chad Territory (General Gov- 
ernment of French West Africa, in litt., November, 1936) . 

In Nigeria "it is pleasant to be able to report that cheetah are 
not nearly as rare as was thought. There are fair numbers in several 
Provinces and they extend nearly as far south as the Benue River." 
("Observer," 1934, p. 54.) 

Senegal Cheetah ; North African Cheetah 


Acinonyx hecki Hilzheimer, Sitz.-ber. Ges. naturf. Freunde Berlin 1913, p. 288, 
fig. 1, 1913. (Based upon a living specimen in the Berlin Zoological Garden, 
said to have come from Senegal.) 

Fios.: Geoffroy and Cuvier, Hist. Nat. Mamm., vol. 3, pi. 145, 1824; Hilz- 
heimer, 1913, p. 287, fig. 1. 

The name A. j. hecki, although based upon a Senegal specimen, 
may be provisionally applied to the Cheetah occurring over the 
greater part of Palaearctic Africa (Mauretania, Morocco, Algeria, 
Tunisia, Cirenaica, and northwestern Egypt). It is a rare form. 

It is described as a small, dainty animal, with a ground color of 
pale reddish ochraceous on the back and sides; spots mostly black, 


but brownish on the cheeks, hind feet, and part of the forefeet; 
under parts white, unspotted except on the lower neck; tail with 
four complete rings and a white tip (Hilzheimer, 1913, pp. 287- 
288). Head and body, 1,150 mm.; tail, 650 mm. (Cabrera, 1932, 
p. 192). 

In Senegal the Cheetah is met with as far south as Podor and 
even near St. Louis (Cligny, 1900, p. 289). 

In Morocco it exists only in the Saharan district, south of the 
Grand Atlas and the Anti-Atlas. Strohl (1923) refers to the capture 
of a dozen specimens in the vicinity of Zenaga [region of Figuig?]. 
According to native report, it is well known, though not very com- 
mon, in the Wadi Draa. Thence it extends across Mauretania to 
Senegal. (Cabrera, 1932, p. 192.) Laurent (1935, p. 350) records 
skins from Tamlelt, Morocco. 

In North Africa the Cheetah is extremely rare, but is still found 
regularly on the Oran-Moroccan High Plateaus. It is also said to 
be distributed here and there in the entire Sahara. (Heim de Balsac, 
1936, pp. 99, 179.) 

In Tunisia this very rare animal exists only in the extreme south, 
in the Grand Erg. Sometimes solitary individuals range toward 
the north; thus some Cheetahs were killed at Fedjej and at El- 
Hamma in 1908 and 1913. The species is hunted by its tracks, 
which are easy to follow on the sand of the dunes. The natives of 
southern Tunisia do not utilize it for hunting, as the Afghans, Arabs, 
and Indians do with the Asiatic Cheetah. (Lavauden, 1932, pp. 7-8.) 

The cause of depletion in Tunisia is the progress of civilization; 
there are no special protective measures. The animal is found ac- 
companying herds of Addax and Loder's Gazelles. (Conservator of 
Forests, Tunis, in Hit., September, 1936.) 

We lack precise information on the range of the Cheetah in the 
Libyan hinterland. Some are found in the southern steppe region of 
Cirenaica. A specimen was killed recently at Bir Scegga, between 
Tobruk and Jarabub. At the time of the Pharaohs the animal was 
employed in the hunting of antelopes. Its skin has slight commercial 
value. (Zammarano, 1930, pp. 13-15.) 

In Egypt the species "is very rare, and found only in the country 
to the west of Alexandria. In 1909 Col. H. C. B. Hopkinson . . . 
saw the tracks of two Chitas that had been stalking gazelle in the 
Mariut district about 40 miles west of Alexandria." In 1910 "a 
Bedawin shot a Chita about 5 miles north-east of Moghara, Mariut 
district .... A few other specimens were shot later." In 1927 
"three live Chita cubs from south-west of Sollum had been received 
recently at the Giza Zoological Gardens." (Flower, 1932, p. 392.) 

The present range of this vanishing species in Egypt is restricted 
to the Western Desert. The cause of depletion is injudicious hunting. 


The skins are sold and the meat is used for food. (Ministry of 
Agriculture and Zoological Garden, Cairo, in litt., January, 1937.) 

Indian Cheetah; Indian Hunting Leopard 

ACINONYX JUBATUS vENATicus (Hamilton Smith) 

F[elis] Venatica Hamilton Smith, in Griffith's Cuvier's Anim. Kingdom, 

vol. 5, p. 166, 1827. ("India.") 
FIGS.: Jour. Bombay Nat. Hist. Soc., vol. 37, no. 4, suppl., pi. 46, 1935 (venati- 

cust); Bodenheimer, 1935, pi. 9 (venations?). 

The Cheetah is nearly extinct in India and has become very rare 
in southwestern Asia generally. 

The general color is pale brownish yellow to bright rufous-fawn 
above and on the sides; almost everywhere with small round black 
spots ; chin and throat buffy white, unspotted ; a black line from the 
anterior corner of the eye to the upper lip, and another less marked 
(or a row of spots in some specimens) from the posterior corner of 
the eye to below the ear; ear black outside, base and margins tawny; 
spots on tail passing toward the end into imperfect rings. Head and 
body, about 4.5 feet; tail, 2.5 feet. (Blanford, 1888, p. 91.) 

The Cheetahs ranging from Baluchistan, Persia, and Iraq to Syria, 
Palestine, and Arabia are here included provisionally with the 
Indian form (A. j. venaticus). 

In India the Cheetah is all but extinct in the wild state. It once ranged 
from the confines of Bengal through the plains of the United Provinces, 
the Punjab and Rajputana, through Central India and the Deccan. ... A 
Cheetah was killed in 1918 and another in 1919 in the Mirzapur District of 
the United Provinces. Five Cheetahs are recorded as having been obtained 
in this Province during the previous twenty-five years. In the Central 
Provinces, the Cheetah appears to have been not uncommon at one time in 
the Berars. Three were shot in the Melghat Forest area in 1890 and one 
in 1894 and one at Wano in 1895. Rumours of their existence in parts of 
Berar, the Seoni Plateau and Saugor still persist. They were apparently once 
common around Hyderabad, Deccan. The only part of the Bombay Presi- 
dency where Cheetahs were known to occur recently is the tract of rugged 
country known as the Tanga in the centre of the province of Kathiawar. In 
1884 it was estimated that there were not more than twenty of these animals 
in this area. A female and four cubs were shot at Rajkot in 1894. (Anonymous, 
1935, p. 147.) 

"In the case of India, the cheetah appears to be verging on ex- 
tinction, if not already extinct, as a wild animal. At all events the 
Mammal Survey of India . . . does not seem to have secured a 
single specimen; and ... it seems that Indian cheetahs are now 
practically unobtainable, and that those used for the chase are 
imported from Africa." (Pocock, 1927, pp. 18-19.) 

"I have heard that Princes and others who want cheetahs for 
hunting purposes now get them from Hyderabad. But the officer 



in charge of the Gwalior shikar department . . . said that 50 or 
60 survived in the state. They are found in Indore also." (Edward 
Thompson, London Times, August 19, 1932?) In Hyderabad State 
"there still remain a few cheetahs" (Salim AH, in Anonymous, 1935, 
p. 231). In Mysore there are several old records (Morris, 1935, p. 
386), but the animal is "probably now extinct" there (Phythian- 
Adams, in Anonymous, 1935, p. 241). Fears are entertained as to 

FIG. 31. Indian Cheetah (Acinonyx jubatus venaticus) 

its survival in the Madras Presidency (M. F. Budge, in litt., Novem- 
ber 12, 1936) . Shooting is completely prohibited (Bombay Natural 
History Society, in litt., December, 1936) . 

"The hunting leopard is certainly found in Persia, but I am unable 
to give any particulars as to its distribution. According to Eichwald 
it does not extend into the countries west of the Caspian, though 
found to the eastward. De Filippi says that it is found in Mazan- 
daran." (Blanford, 1876, p. 35.) To this statement 0. St. John adds 
that the species "is not used at present for sporting purposes in 
Persia." [The form of northern Persia may be A. j. raddei.] 

"As regards its survival in Persia, Arabia and Palestine, I have 
no information beyond the inference to be drawn from the infre- 
quency with which it is mentioned" (Pocock, 1927, p. 18) . 

"In South-Western Asia its range is believed to reach from the 


frontiers of Sind through parts of Afghanistan, Baluchistan and 
Persia and Mesopotamia to Syria and Palestine .... To what 
extent it survives in these Asiatic countries is not known." (Anony- 
mous, 1935, p. 147.) 

In 1925 a cub was secured at Jumaimah, Muntafq, Iraq, and in 
1928 two cubs were taken near Busiya on the Shamiyah Desert. 
None of the local Arabs had seen a Cheetah before. (Corkill, 1929, 
pp. 700-702.) 

Danford and Alston (1880, pp. 52-53) report on the Cheetah in 
Syria as follows: "A skin of the Cheetah was presented to Danford 
at Biledjik, on the Euphrates, by his host Sheik Mustapha, who 
stated that the animal had been killed among the rocks near Sevi, 
a small village about five hours down the river on the Mesopotamian 
side ; it was the only specimen which he had ever seen. This Society 
[the Zoological Society of London] has received more than one 
specimen from Syria, and it is not improbable that the species may 
be found in some parts of Asia Minor proper." 

Tristram wrote in 1884 (p. 19) of this species in Palestine: "This 
graceful Leopard is scarce, but still haunts the wooded hills of Galilee 
and the neighbourhood of Tabor. East of Jordan it is far more com- 
mon, and is much valued by the Arabs." 

The Cheetah has now become very rare in Palestine. Yet it is 
still pretty common in the southern steppe. Its use for the chase is 
now quite outmoded. (Aharoni, 1930, p. 332.) It "still lives in the 
Negeb, in Transjordania and rare specimens also persist in the Pales- 
tinian mountains. The author saw many skins, sold by Beduins 
from Beersheba." (Bodenheimer, 1935, p. 105.) More recently Pro- 
fessor Bodenheimer writes (in litt., March, 1937) that the animal 
is now on the verge of extinction or extinct and that nothing can be 
done to preserve it in Palestine, but that perhaps there is still a 
chance to do so in Transjordania. 

In 1909 Carruthers (1935, pp. 60, 70) found Cheetah tracks on the 
north side of the Jabal Tubaiq, Arabia, approximately 150 miles 
east of the head of the Gulf of Akaba. 

Turkestan Cheetah 


Acinonyx raddei Hilzheimer, Sitz.-ber. Ges. naturf. Freunde Berlin 1913, no. 5, 
p. 291, 1913. (Based upon a specimen purchased in Merv, Russian 
Turkestan (Turcoman S. S. R.).) 

This Cheetah seems to occur in very small numbers in the southern 
parts of Russian Turkestan. The animals of northern Persia and 
northern Afghanistan may belong to the same form. 


It is distinguished by its extremely thick, long fur; ground color 
light brownish gray ; very large spots reaching to the toes ; tail long- 
haired and very bushy, with five half -rings (one perhaps a complete 
ring) at the end (Hilzheimer, 1913, p. 291). (Cf. Satunin, 1909, 
pp. 254-256.) 

"Only in the western portion of Turkestan have I met with this 
species, and even there only on the low plains" (Severtzoff, 1876, 
p. 49). 

This is doubtless the commonest of the large cats in Transcaspia. 
It is distributed through the whole region on the lowlands, along 
the river courses, and on the mountains. Each year the Turkomans 
bring young Cheetahs for sale into the cities and military posts. 
Training the animals for the chase is unknown to them. (Radde and 
Walter, 1889, p. 1012.) 

The following data are from Ognev (1935, pp. 313-314) : The 
Cheetah is found from time to time as far north as the Mangyshlak 
Peninsula (Karelin, 1883). It inhabits the Kara Tau, the western 
spurs of the Tian Shan, the lower Syr Darya, the Zarafshan Valley, 
and the steppes between Zarafshan, Syr Darya, and Kizil Kum, 
reaching an elevation of 600-1,000 feet (Severtzov, 1873). It also 
occurs on the Amu Darya (Zarudny, 1915) and in Tajikistan. It is 
particularly numerous along the Murgab, Tejend, and Sumbar 
Rivers (Bilkewicz, 1918). The Caucasian Museum has specimens 
from Merv, Kizil Arvat, and the Kopet Dagh. The Cheetah is also 
recorded from Mazanderan, northern Persia (De Filippi) . 

About 1884 two cubs were obtained in northeastern Persia near 
the Turbat-shaikh-jami River, a tributary of the Hari Rud (Ait- 
chison, 1889, pp. 56-57) . 

The Cheetah is observed irregularly and in very small numbers 
in Turkestan on the frontiers of Persia and Afghanistan. Some are 
killed, but not every year. (W. G. Heptner, in litt., December, 1936.) 

King Cheetah; Cooper's Cheetah 


Acinonyx rex Pocock, Abstr. Proc. Zool. Soc. London, no. 283, p. 18, Mar. 1, 
1927 ("Umoukwe [=Umvukwe] Range, N.W. of Salisbury, [Southern] 
Rhodesia"); Proc. Zool. Soc. London 1927, pt. 1, p. 250, pi. 1, April 6, 

FIGS.: Pocock, 1927a, pi. 1, and 19276, frontisp.; Dollman, 1929a, p. 3, fig.; 
Maydon, 1932, pi. 103. 

The range as well as the numbers of the King Cheetah are dis- 
tinctly limited, and special protective measures seem to be neces- 
sary in order to insure its survival. 

It is similar in size and proportions to the Common Cheetah (A. 
jubatus) but distinguished from it by a bold pattern of black 


stripes and blotches, which are longitudinal on the dorsal area and 
oblique or longitudinal on the flanks; legs blotched and spotted to 
the feet; basal half of tail with two longitudinal stripes, distal half 
with irregular transverse stripes; ground color mostly cream-buff; 
belly white. Skin measurements: head and body, 4 feet 2 inches 
to 4 feet 5 inches; tail, 2 feet 6 inches. (Pocock, 1927a, pp. 250- 
252.) The King Cheetah is by some considered as a color mutation 
of the Common Cheetah (G. M. Allen, MS.). 

FIG. 32. King Cheetah (Acinonyx rex). After Pocock, 1927. 

The species is known only from Southern Rhodesia. Its range 
may lie wholly within the northern division of the Southeastern 
Veldt District of Bowen (1933, pp. 256, 260) . 

This superb new Cheetah was brought to scientific attention in 
1926, by Major A. L. Cooper, who sent to the British Museum the 
skin of an animal trapped by natives in the Umvukwe Range 
(Pocock, 1927a, p. 245). 

Cooper (in Maydon, 1932, pp. 335-336) gives the following ac- 

"That this animal was known for some time past is borne out 
by the fact that, twenty years ago, mention used to be made round 
camp fires by natives of a beast that was neither Lion, Leopard, 
nor Cheetah, and ... I believe was referred to as the 'Mazoe 
Leopard.' It was apparently commoner in those days than it is now." 

The skin now in the Salisbury Museum was purchased from 
natives, who stated that they had killed the animal in the Macheke 
district. There were four or five in the troop. 

H. M. G. Jackson reported a similar skin at the American Mission 


at Utambara. "It was also an old native police sergeant of his, 
who, when shown the skin, said he knew the animal, told us its 
native name, and informed us of its habits, namely, that it is ex- 
tremely shy, never attacked domestic animals except possibly a 
young kid, and, when chased by dogs, never took to a tree as a 
Cheetah occasionally does. . . . 

"It was found that Mr. Watters, Native Commissioner at Bitika, 
possessed two such skins. . . . These were presumably obtained in 
his district. Apart from these two, I found Mr. Lacey of Salisbury 
also had a specimen, . . . killed some twenty miles south of Salis- 
bury. . . . 

"This is the history of the discovery, if it can be described as 

Pocock reports (1927a, p. 246) that the animal whose skin was kept 
at the Utambara Mission "was shot in the Melsetter District close 
to the Portuguese Border. . . . The natives were not at all afraid 
of it as they were of leopards, and would attack it armed only with 
assegais." He says also (19276, p. 19) : 

In the interests of the preservation of the new species of cheetah the 
following probabilities cannot be too strongly insisted upon. All the avail- 
able evidence suggests that the animal has a restricted range and is nowhere 
plentiful. Its distributional area is within reach of Salisbury, an easily 
accessible centre; and the publicity now given to the existence of so hand- 
some an animal will surely be taken advantage of by sportsmen and traders. 
All the big museums in the world will be eager for its skin, and every 
zoological garden will want live specimens for exhibition. It will, therefore, 
command a high price, whether alive or dead, and the result will be per- 
secution by hunters and trappers on such a scale as to threaten its extinction 
unless the authorities in Rhodesia at once take such steps as may be neces- 
sary to protect it. 

There seem to be no nature reserves within the known range of 
the King Cheetah. 

Barbary Lion. Le Lion de Barberie (Fr.). El Leon 
berberisco (Span.) 

LEO LEO LEO (Linnaeus) 

Felis leo Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 41, 1758. ("Africa"; type locality 
subsequently restricted to Constantine, Algeria, by J. A. Allen, Bull. Am. 
Mus. Nat. Hist., vol. 47, p. 222, 1924.) 

FIGS.: Reichenbach, 1836, figs. 1-2; Geoffroy and Cuvier, Hist. Nat. Mammif., 
vol. 1, pis. 114-115, 1824. 

The Barbary Lion survived well into the twentieth century but 
is now extinct. 

"Very large, dusky ochery, with the mane very thick and long, 
extending to the middle of the back; and a thick and heavy mane 


on the under-parts. In the female the inside of the fore-legs is 
white." (Lydekker, 1908, p. 416.) The female is paler and smaller 
than the male (Cabrera, 1932, p. 181). 

This extinct race ranged from Tripoli through Tunisia, Algeria, 
and Morocco. Apparently there are no definite records of Lions 
within the present confines of Egypt during historic times. The 
Lion of Senegal is described as a distinct race (Leo leo senegalensis 
(J. N. von Meyer)), although a single race of Cheetah (Adnonyx 
jubatus hecki Hilzheimer) is accredited to both Senegal and the 
Barbary States. 

Tripoli. "About two hundred years ago the lion was found quite 
commonly in Tunisia. About the same time*, so far as records go, the 
last lion was killed in the adjoining Pashalik of Tripoli, where the 
animal now seems to be entirely extinct." (Johnston, in Bryden, 
1899, p. 564.) 

Tunisia. "Down to the time of the French invasion of Tunis, 
in 1881, lions were still found in the extreme north-western part of 
the Regency, close to the Algerian frontier. . . . 

"What has brought about the extinction of this animal is less 
the persistent attacks of French or Arab sportsmen than the opening 
up of the forests and the settling down of the people since the French 
occupation. The herds are now so carefully tended that the lion has 
little or no chance of feeding on them, while the Barbary stag and 
the gazelles have in that region become very scarce." (Johnston, in 
Bryden, 1899, pp. 562-564.) 

The last Lion of Tunisia was killed in 1891 at Babouch, between 
Tabarka and Am-Draham. The species was common up to the time 
of the French occupation. Doubtless owing to troop movements, it 
then retired to the most remote massifs. It could not survive in 
contact with civilization. (Lavauden, 1932, pp. 5-6.) 

Two specimens in the Ley den Museum (one killed in Tunisia in 
1823, the other in "North Africa" about the same period) are prob- 
ably the only wild-killed Barbary Lions that are preserved in any 
museum and can be studied at present (Cabrera, 1932, pp. 182-183). 

Algeria. Pease (in Bryden, 1899, pp. 564-566) gives the following 

The North African lion was in bygone ages undoubtedly very numerous. . . . 

The Algerian lion has become so rare that it may be said to be nearing 
extinction. ... It lingers only in the country that might almost be 
described as the Mediterranean littoral zone, though an occasional lion is 
still shot or tracked in the interior, as far inland as the district of Soukarras, 
and certain places in the Aures. [During 1892-95] I do not remember hearing 
of more than three or four being obtained in the whole province of Con- 
stantine. In the provinces of Algiers and Oran they may be said to be ex- 
tinct. So long ago as 1862 General Marguerite wrote that ... in the province 
of Algiers . . . the average number killed did not exceed three or four a 


year. . . . General Marguerite relates, that during his eleven years the 
Beni-Mahrez, a tribe not numbering more than 100 tents, lost on average 
annually, 3 horses, 25 cattle, and 75 sheep from the depredations of lions 
and panthers .... Before the French came, the Turks had encouraged the 
Arabs to destroy them by freeing the two great lion-hunting tribes, the 
Ouled Meloul and Ouled Cessi, from all taxes and paying liberally for their 
skins. The French gave only 50 francs for a skin. 

Between 1873 and 1883 the process of extinction is measured in Govern- 
ment returns. The numbers killed for the whole of Algeria were, in the last 
six years of this period, 1878, 28; 1879, 22; 1880, 16; 1881, 6; 1882, 4; 1883, 3; 
(1884, 1) ; and for the decade- 
Province of Algeria 29 

" " Constantine 173 

" " Oran 


There are a few lions still left in the Province de Constantine, in the thick 
forests between Soukarras and La Calle. 

According to Johnston (in Bryden, 1899, p. 564), "Lions still 
linger here and there in South-East and South-West Algeria." 

On the other hand, Lavauden, an eminent authority on the North 
African fauna, fixes (1932, p. 6) the date of the Lion's disappearance 
in Algeria at about 1891, when the last one was killed in the region 
of Souk-Ahras. 

Morocco. To judge by the literature of several centuries ago, 
Morocco was then a veritable country of Lions. At the middle of 
the seventeenth century they still abounded on the Mediterranean 
coast. Toward the end of the eighteenth century the range extended 
to Cape Nun, Ifni. By the middle of the nineteenth century the Lion 
had retreated from the entire Mediterranean littoral. According to 
old native hunters, about 1880 not a Lion remained north of the 
low Bu Regreg and Taza Pass. Some years later many forests of the 
Middle Atlas served as a refuge for bandits, and this fact, together 
with the civil wars of those times, contributed to the disappearance 
of the Lions. Even in 1901 Lions were said to be frequent visitors 
to the forest of Budaa, near Azru. According to the ex-Sultan Muley 
Hafid, there remained in Morocco about 1911 only a few Lions, 
which lived in the forests of the Zaian and the Beni Mguild. Ap- 
parently they survived at least to 1922 in the Middle Atlas, and 
it is probable that they inhabited the Grand Atlas likewise till a 
comparatively recent date. They inhabited especially the wooded 

The Lion figures largely in the folklore of Morocco. 

Its rapid disappearance from this country constitutes a very 
curious problem. Unlike the Cape Province and Algiers, Morocco 
remained wild and uncivilized up to a quarter of a century ago. 
Its inhabitants are far from being a hunting people, and few Euro- 


peans have ventured there owing to the lack of security. (Cabrera, 
1932, pp. 186-190.) 

Heim de Balsac (1936, p. 98) places the disappearance of the 
species in Morocco in the decade 1900-1910 a considerably earlier 
date than Cabrera's. 

Utilization by the Romans. "There is ... little doubt that the 
Romans drew their chief supply of lions for the arena and gladia- 
torial combats from Mauretania and Numidia." Pliny speaks of 
hundreds at a time being shown by Pompey and Caesar in the 
Roman arena. (Pease, in Bryden, 1899, p. 564.) This bespeaks a 
great abundance of Lions in North Africa at that period. 

European Lion 

LEO LEO subsp. 

Of the Lion that still existed in Greece in classical times, no 
remains seem to have been found. If suitable material were avail- 
able, the modern systematist would probably find means of dis- 
tinguishing it from any living Lion as well as from its Pleistocene 
ancestor (Leo spelaeus) that once roamed over a large part of 
Europe. Up to the present time, however, it apparently has not 
received even a subspecific designation. 

Meyer (1903, pp. 65-73) has provided a useful summary of our 
knowledge of the European Lion. Herodotus (ca. 484-430 B. C.) 
reports many Lions between the Achelous River in Acarnania and 
the Nestus River in Abdera (Thrace) and states that during 
Xerxes's march through Macedonia (480 B. C.) Lions killed some 
of his baggage camels. Aristotle (384-322 B. C.) assigns the same 
range to the Lion, but speaks of it as rare. By A. D. 80-100 it was 
considered entirely exterminated in Europe, as a result of a gradual 
retreat before man and his culture. 

"The Greek name for the lion is very ancient, and this suggests 
that it refers to an animal indigenous to the country. Although the 
evidence is not decisive, it seems probable that lions did exist in 
Greece at the time of Herodotus; and it is quite possible that the 
representation of a lion-chase incised on a Mycenean dagger may 
have been taken from life." (Flower and Lydekker, 1911, vol. 16, 
p. 737.) 

Evidently Elliot regarded this Lion as identical with the pre- 
historic Cave Lion (Leo spelaeus). He writes (1883, text to pi. 1) : 
"The Cave-Lion disappeared from Britain towards the close of the 
Postglacial period, and is considered to have retreated gradually 
from Europe and become extinct between 340 B. C. and A. D. 100. 
The cause of this disappearance, according to Dawkins, was the 
warfare carried on against it by the people of those periods, as 


exhibited by the leonine remains found in the ancient dwelling 
places of the Postglacial men in Aurignac and La Madeleine. This 
is probably a correct supposition; for neither was the temperature 
unsuited for its continued existence nor had the supply of food 

Asiatic Lion; Indian Lion 


Felis leo persicus Meyer, Dissertatio inauguralis anatomico-medica de genere 

Felium [Vienna], p. 6, 1826. (Persia.) 

SYNONYMS: Felis leo goojratensis Smee (1833); Leo asiaticus Jardine (1834). 
FIGS.: Trans. Zool. Soc. London, vol. 1, pi. 24, 1834; Jardine's Nat. Libr., 

vol. 15, Mammalia, pi. 11, 1842; Elliot, 1883, pi. 1, upper right-hand fig.; 

Pocock, 1930, pis. 1-3. 

This Lion, once widely distributed from Asia Minor, Palestine, 
and Arabia to Persia and India, is now almost or entirely reduced 
to a small remnant in the Province of Kathiawar, in western India. 
There may be also a few solitary survivors in Persia and Iraq, but 
this is doubtful. 

"On the average, the Indian lion has a scantier mane than the 
African and, curiously enough, ... a fuller coat, a longer tassel 
of hair at the end of the tail, a more pronounced tuft of hair on 
the elbow joints and a fuller fringe of hairs on the belly. In size, 
there is little to choose between the two. . . . The largest recorded 
measurement of an Indian lion is 9 ft. 7 in., of an African lion 
10 ft. 7 in." (Anonymous, 1935, p. 123.) 

Asia Minor. In 1878 or 1879 Sheik Mustapha informed Danford 
"that five years ago a Lion appeared near Biledjik [on the Euphrates 
toward the Syrian border], and after destroying many horses was 
done to death" (Danford and Alston, 1880, p. 53). This is the only 
definite locality record I have found for Asia Minor. 

Syria, Palestine, and Arabia. "The Lion has long been extinct 
in Palestine, and among the inhabitants there is no tradition of its 
existence. Yet of its former abundance there can be no question. 
It is mentioned about 130 times in Scripture .... Within the 
historic period it was common in Syria, Asia Minor, and Greece. 
... It seems to have disappeared altogether from Palestine about 
the time of the Crusades, the last mention of it being by writers of 
the twelfth century, when it still existed near Samaria. ... It can 
scarcely be said now to exist in Asia west of the Euphrates, unless 
in Arabia, the latest trace being that a few years ago the carcase of 
one was brought into Damascus. . . . The Arabs state it is found 
in Arabia." (Tristram, 1884, p. 17.) 

According to the Old Testament, the Lion was found in Lebanon 


and on the Jordan. The ancient writers (Xenophon, Aristotle, 
Strabo, Pliny, etc.) speak of lion hunts in Syria and also in Arabia. 
(Meyer, 1903, p. 71.) 

"To-day, the nearest [to Palestine] wild habitats of the lion are 
the jungles of the Upper Euphrates and several Arabian oases. But 
even in those places it must be on the verge of extinction." (Boden- 
heimer, 1935, p. 114.) 

Iraq (Mesopotamia) and Persia. In the early part of the last 
century Lions were noted fairly commonly along the Euphrates and 
the Tigris. The explorer Layard hunted them with the Bakhtiyari 
chiefs in Arabistan, whose sheep and oxen suffered from the Lions' 
depredations. By the middle of the century Layard reported the 
species as then found rarely on the Tigris as far north as Mosul, 
but frequently below Bagdad. He adds: "On the Euphrates it has 
been seen, I believe, almost as high as Bir .... On the [Jebel?] 
Sinjar and on the banks of the Khabour [in the northeast of the 
present Syria], they are frequently caught by Arabs. They abound 
in Khuzistan [western Persia]." (Kinnear, 1920, pp. 33-35.) 

By 1891, according to Sir Alfred Pease (Book of the Lion), the 
"lion is no longer found in Asia Minor, but exists in Mesopotamia 
and Arabistan, between Poelis, west of Aleppo, and Deyr [in the 
present Syria] , and in the Euphrates valley ... ; it is also found in 
the lower part of the Karun river but is nowhere plentiful." (Kin- 
near, 1920, p. 36.) 

Blanford writes (1876, p. 29) : "The lion at the present day is 
found in Mesopotamia, on the west flanks of the Zagros mountains 
east of the Tigris valley, and in the wooded ranges south and south- 
east of Shiraz. It nowhere exists on the table land of Persia." To 
this 0. St. John adds (in Blanford, 1876, pp. 30-31) : "Lions, which 
are very numerous in the reedy swamps bordering the Tigris and 
Euphrates, are found also in the plains of Susiana, the modern 
Khuzistan, and extend into the mountain country south of Shiraz 
as far east as longitude 53." Acorns of an oak (Quercus aegilopi- 
folia) "feed the wild pigs whose- presence tempts the lion into the 
mountains of Pars. . . . The little valley of Dashtiarjan, thirty- 
five miles west of Shiraz, is notorious for the number of lions found 
in its vicinity. . . . Dashtiarjan is ... a perfect paradise for 
swine, ... so that the lions have plenty to eat .... Every year 
some four or five adult lions are killed in Dashtiarjan or the neigh- 
bourhood, and a few cubs are brought in to Shiraz for sale." 

Edward Thompson (London Times, August 19, 1932?) gives the 
following reports for Mesopotamia: a Lioness and cubs seen by an 
Indian trooper near Ahwaz in 1917; a Lion cub brought through an 
Arab village near Sanniyat in 1916; and one shot in the Wadi 


marshes a year later. He also mentions a report received by the 
Bombay Natural History Society of the continued existence of Lions 
in the Pusht-i-Kuh range of western Persia. 

"There are Persian Lions, and the last time a pair of them was 
seen in the South of Persia by French and English Engineers (in 
1928) .... The animals were carefully watched for several hours 
and were seen by hundreds of people. ... I understand that the 
Persian ruler takes keen interest in their preservation and they 
are not allowed to be shot." (Hasan Abid Jafry, in Hit., August 
17, 1933.) 

The Persian Lion is a thing of the past. Firearms, whose use 
increased during the World War, were more dangerous to the Lion 
in Persia than in Africa. In 1923 the last of its kind was killed 
south of Shiraz. Yet the people still express belief in the existence 
of Lions. In the swamp and reed areas of the Euphrates and the 
Tigris I have been able to find no more trace of the Lion. The opera- 
tions here during the World War paved the way for its extinction. 
Skins of Persian Lions are still found in some mosques. (Becker, 
1934, pp. 439-440.) 

The Lion may survive in the wilder mountains of Luristan and 
Khuzistan in southwestern Persia (Bombay Natural History Society, 
in litt., December, 1936) . 

"The Syrians frequently used the lion motif as a frieze decora- 
tion, and at Persepolis, thirty miles northeast of Shiraz, where the 
magnificent ruins of the palace of Darius the Great may still be 
seen, the lion as a decorative architectural motif was constantly used. 
In the embrasures of some of the great doors of Persepolis the winged 
lions were magnificently carved." (Vernay, 1930, p. 82.) 

Afghanistan and Baluchistan. "There is no evidence to show 
that the lion inhabited Afghanistan or Baluchistan within historic 
times" (Kinnear, 1920, p. 37) . 

"I was told, while in Duzbad, the frontier town on the Baluch- 
Persian border, that the lion existed in Afghanistan seventy-five 
years ago. This is mere heresay, but it sounds quite reasonable." 
(Vernay, 1930, pp. 82-83.) 

In 1935 Admiral Philip Dumas reported seeing a Lion at close 
range near the Bolan Pass, south of Quetta in Baluchistan (Jour. 
Bombay Nat. Hist. Soc., vol. 38, no. 2, pp. 381-382, 1935). 

India. "Within the present [19th] century, distributed over 
much of Central, West, and North-west India; but now confined 
in that country to the peninsula of Guzrat, unless a last remnant 
still maintains a lingering existence in the jungles bordering the 
Sind River in Bundelkund, which I now consider doubtful" (Blyth, 
1863, p. 182). 


Kinnear (1920, pp. 37-39) writes: 

[The Lion] was formerly found in Sind, Bahawalpur and the Punjab, 
becoming extinct round Hariana, in the latter province, in 1842. It was 
however extinct in Sind before that date and the last on record was shot 
near Kot Deji in 1810. Exactly how far eastwards the lion was a regular 
inhabitant we do not know, though there is a statement of one being killed 
in the Palamaw district, Behar and Orissa, in 1814, but whether this was 
merely a straggler or not, there is no evidence to show. The southernmost 
limit appears to have been the Narbada. In 1832 one was killed at Baroda, 
while further north it was comparatively common round Ahmedabad in 
1836. Central India in these early days was one of the strongholds of the 
lion and to give an idea of its numbers we may mention that Lydekker was 
informed that during the Mutiny, Colonel George Acland Smith killed up- 
wards of 300 Indian lions and out of this number 50 were accounted for in the 
Delhi district ! 

The occurrence of the lion in Cutch is doubtfully recorded. The lion 
probably was found in Cutch at one time but the records are not satisfactory. 

Dates of extermination in other parts of India, according to Kin- 
near, are: Damoh district (Saugor and Narbada territories), 1847- 
48; Rewah (between Allahabad and Jubbulpore), 1866; Goona, 
1873; Abu and Jodhpur (Rajputana), 1872; Deesa (Guzerat), 1878; 
Palanpur (Guzerat), 1880. 

A map, showing dates of extermination of the Lion in various 
localities in India, is given by Pocock (1930, facing p. 661). 

"A small number [of African Lions] were imported into Gwalior" 
about 1890-1900, "but after a few years they became a pest, killing 
not only the cattle of the natives, but also the natives themselves, 
so that the African lions were all eventually shot out. Also, the 
tigers of Gwalior are famous, and as tigers will not permit lions to 
remain in their territory, they must have helped to kill off the lions." 
(Vernay, 1930, pp. 81-82.) 

"In India the lion is verging on extinction. There are probably a 
very few still living in the wild tract known as the Gir in Kattywar, 
and a few more in the wildest parts of Rajputana, especially South- 
ern Jodhpur, in Oodeypur, and around Mount Abu." (Blanford, 
1888, p. 57.) 

"In 1893 ... a rough census was taken [in the Gir Forest], 
and the number remaining was estimated at twenty-six, which sub- 
sequent estimate raised to thirty-one. . . . There are now esti- 
mated to be only twenty lions remaining in the Gir, of which eight 
are cubs. (Lydekker, 1900, pp. 270-271, quoting from The Asian 
newspaper of June 19, 1900.) 

"It is only in the Province of Kathiawar, a small peninsula north- 
west of Bombay, that the true Asiatic lion can still be found. Even 
there it exists only in the Gir Forest, an area of four hundred square 
miles in the State of Janagadh. . . . 


"It is only a question of time before the lion will disappear even 
from this district, although at present it is closely protected. The 
number, which is roughly estimated at 200, is not increasing. The 
inevitable diminution of the forest, in spite of the restrictions against 
cutting, and the possibility of disease owing to the confined area, 
mean ultimate extermination." (Vernay, 1930, p. 81.) 

Economics and conservation. The Lions "commit considerable 
havoc amongst the cattle, which are brought into the Gir for grazing 
purposes during the greater part of the year, besides helping them- 
selves liberally to the sambar, nilgai, spotted deer, and pig with 
which the forest abounds. ... A large number of lions are kept 
in captivity in the State gardens at Junagarh, where they breed 
very freely." (L. L. Fenton, in Lydekker, 1900, pp. 410-412.) 

"It is reasonable to suppose that the factors which exterminated 
it in Europe, Asia Minor and Syria and have brought it to the verge 
of extinction in Mesopotamia and Persia, even if they have not 
already achieved that end, were the same as the factors which 
exterminated it over almost the whole of the area it occupied in 
India. In my opinion there is no reasonable doubt that the main, if 
not the sole, factor in the case of Europe and southwestern Asia 
was man. At all events it was most emphatically not the tiger. . . . 

"It is not unlikely, in my opinion, that the Kathiawar stock is 
deteriorating in size from inbreeding." (Pocock, 1930, pp. 641, 665.) 

"I ... hear that the status of the Indian lion, as far as preser- 
vation and numbers are concerned, is most satisfactory. They have 
of late been overflowing from their original reservations in the 
Gir Forests of Kathiawar and Junagadh State, and have made 
themselves unpopular by cattle killing. Their numbers are esti- 
mated to be not less than 150. 

"It must be remembered that there is no wild life in their present 
habitat on which they can prey, and they live almost entirely on 
village cattle. There are, of course, far more cattle there, as in other 
parts of India, than are economically desirable ; but ... if the local 
native rulers were to withdraw their protection, the lion would 
speedily disappear. There is, however, little danger of this happen- 
ing, and the villagers at present cooperate loyally in the protection 
of these animals; even to the extent, in a recent case, of pulling a 
lion out of a well into which he had fallen, with no little risk to 
themselves." (C. H. Stockley, in Hit., May 29, 1933.) 

"Even in the Province of Kathiawar, where tigers do not exist 
and where no struggle for supremacy between these two giants of the 
tribe could have taken place, the lion was slowly driven from the 
Barda and Aleche hills, from parts of Dhrangadra and Jasdan as a 
result of human settlement and the progress of cultivation. 


"The number of lions in the Gir is computed to be well below a 
hundred." (Anonymous, 1935, p. 125.) 

Cadell (1935, pp. 165-166) writes as follows: 

The animals are easily enticed across the boundary [of Junagadh State] 
by a succession of tie-ups. ... To our certain knowledge . . . twenty- two 
animals have been so slain within the three seasons ending in 1934. . . . 

If every year the State has the very real honour and pleasure of enter- 
taining distinguished guests for a lion shoot, it is a distinction which costs 
a good many thousands of rupees. There is also the steady annual cost of 
the sums paid in compensation to villagers and herdsmen whose cattle have 
been killed by lions. . . . 

There were supposed to be less than a dozen [lions] in 1880 .... As a 
result of the strict preservation during the [British] Administration [from 
1911 to 1920] the number was believed to have increased to fifty .... It 
has since been stated . . . that there are now two hundred lions. . . . My 
own opinion ... is that there are not much more than 75 to 80. ... 

The pressure on Junagadh of suggestions for invitations to shoot lions 
is ... increasing year by year .... Unless an agreement is reached [to 
limit the number shot in one year to some such figure as five or six], and is 
faithfully observed, the danger of the disappearance of the lion from the 
fauna of India, and consequently from its last home in Asia, is obvious. 

Cape Lion. Leeuw (Boer) 


Leo melanochaitus Hamilton Smith, Jardine's Naturalist's Library, vol. 15, 
Introd. to Mammalia, p. 177, 1846. ("Cape of Good Hope.") 

FIGS.: Griffith's Anim. Kingdom, vol. 2, pi. facing p. 428, 1827; C. H. Smith, 
1846, pi. 10; Harris, 1840, pi. 29; Pocock, 1931, p. 208, lower fig. 

The Cape Lion was the first of the African subspecies of Leo leo 
to become extinct. The last record for the Cape Province is ap- 
parently 1858; for Natal, 1865. 

"The species is of the largest size, with a bull dog head ; . . . large 
pointed ears edged with black; a great mane of the same colour 
extending beyond the shoulders; a fringe of black hair under the 
belly; a very stout tail, and the structure in general proportions 
lower than in other Lions" (C. H. Smith, 1846, p. 177). 

Pocock (1931, p. 208) writes as follows concerning a mounted 
specimen in the Junior United Service Club, London, which is "said 
to have been killed near the Orange River about 1830, probably 
. . . near Colesberg": 

The mane is not only remarkable for its luxuriance, length and extension 
over the shoulder, but also for its blackness. It is indeed wholly black except 
for the tawny fringe round the face and a certain amount of the same pale 
hue low down on the shoulder. 

The elbow-tuft and tail-tuft are likewise big and black ; but the belly fringe, 
long and thick behind, becomes gradually shorter and thinner and gradually 
disappears in front of the chest. 

The interest of this lion lies in its being, so far as I am aware, the only 


representative, in this country at all events, of the now extinct race of 
splendid liong which formerly inhabited Cape Colony. . . . 

The former range and the date of the extermination of the handsome South 
African race are alike unknown. 

Pocock adds that Smith's type specimen appears to have been a 
rickety captive. 

Roberts (1929, p. 92) quotes from Paterson (1789) the following 
measurements of a lioness from the southern part of Cape Province : 
total length, 8 feet 9^ inches; tail, 3 feet; "height before," 3 feet 
8 inches. 

Owing to lack of material, the exact limits of the range of the 
Cape Lion will never be known. For present purposes the Cape 
Province and Natal will be considered to comprise the former range. 

"Civilization's steady march in South Africa during the past 
twenty years has considerably limited the range of the lion. The 
vast herds of game upon which he depended for food being swept 
away, he has been forced to retire into remoter regions. From much 
of the South Africa of Gordon Gumming he has vanished com- 
pletely and forever." (Kirby, in Bryden, 1899, p. 549.) 

"With regard to past times Kolben (1731), states that lions 
were not uncommon near Cape Town as late as 1707, Sparrman 
(1785), Paterson (1790), Thunberg (1795), and Barrow (1801), all 
met with these animals as soon as they got away from the imme- 
diate neighbourhood of Cape Town especially on the karoo and in 
Uitenhage. The last record I have met with of the occurrence of a 
lion south of the Orange River is of one killed with assegais near 
Commetjes Post on the eastern frontier in 1842, as noted by Hall. 
General Bisset shot a lion in Natal in 1865, which is probably the 
last record for that Colony." (W. L. Sclater, 1900, vol. 1, p. 31.) 

"Their food . . . consists of the larger game, mainly antelopes 
of all kinds, but also includes zebras, giraffes, and buffaloes. They 
will kill the donkeys and cattle belonging to prospecting and hunt- 
ing parties, and will raid Kaffir kraals when driven to it by hunger. 
Man-eating lions are generally old animals with bad teeth." (Haag- 
ner, 1920, pp. 69-70.) 

"It is stated that a lion was shot on the Ingonyama Tributary 
of the Tsomo River, Transkei, in 1858. One was reported from Port 
Alfred in 1846, and one was killed by shot from a spring gun on the 
farm Lombards Post near Southwell, near Bathurst, about 1850." 
(Hewitt, as quoted by Shortridge, 1934, vol. 1, p. 80.) 

The nearest areas to the former range of the Cape Lion, that are 
still inhabited by some form of the species, are South-West Africa, 
the Kalahari, and eastern Transvaal. The last-mentioned area 
(especially the Kruger National Park) is the home of the Sabi 
Lion (Leo leo krugeri Roberts) . 


Manchurian Tiger; Siberian Tiger; Amur Tiger; 
Mongolian Tiger 


Tigris longipilis Fitzinger, Sitz.-ber. Akad. Wiss. [Wien], math.-nat. Cl., 
vol. 58, pt. 1, p. 455, 1868. ("Korea and Japan through northern China, 
Manchuria, Mongolia, and Dzungaria north to southern Siberia, and west 
through northern Tatary, Bokhara, and northern Persia to Mount Ararat 
in western Armenia"; type locality restricted by Lydekker (1901, p. 288) 
to "Amurland." Cf. Harper (1940, p. 194).) 

SYNONYM: Felis tigris var. amurensis Dode (1871). 

FIGS.: Pocock, 1929, pi. 4, pi. F (upper fig.); Morden, 1930, p. 548, fig.; 
Pocock, 1937, p. 770, fig. 

This Tiger, whose principal range is in northern Manchuria and 
southeastern Siberia, seems to be declining rather rapidly in numbers. 

It is somewhat larger and has a longer and thicker coat than 
the Bengal Tiger; ground color paler; stripes less pronounced and 
tending to become brown on the flanks. It is said to reach a length 
of 13-14 feet. 

Siberia. The following information is from Ognev (1935, pp. 
292-293). Radde (1862) found the species along the Argun River 
near Ust Strielka and near Nerchinskiy Zavod. Baikov (1925) 
places its northern limits at the Shilka and the lower Zeya and 
Bureya Rivers. It is numerous in certain parts of the southern 
Ussuri district. A specimen in the Zoological Museum of the Uni- 
versity of Moscow was said to have been taken in 1828 near Ba- 
lagansk, west of Lake Baikal (Severtzov, 1855). (This record can 
not be definitely allocated as to subspecies.) 

Ford Barclay (1915, pp. 225-228) gives the following account: 

Careful inquiries made in the summer of 1899 along the present route of 
the Siberian Railway, as far as Chita and Niertschinsk on the Amur and 
thence east along that river as far as Khabarovsk, elicited practically no 
information .... 

At Khabarovsk . . . plenty of information was forthcoming, and many tales 
were floating about of the depredations of these animals during the winter 
in close proximity to, and even in one case within, the town itself. . . . The 
best ground was reported to be in the neighbourhood of Irma, ... a little 
more than half-way to Vladivostok, [where large numbers of Wild Pigs 
attracted the Tigers.] 

At Irma I learnt that a number of skins were undoubtedly brought in every 
winter, but it was believed that in most cases their wearers had been accounted 
for by poison. . . . 

In 1899 it was still not uncommon to find fresh footprints of tiger on a 
winter's morning in any of the outlying streets of Vladivostok .... 

In the mountainous district between Harbin and Vladivostok a certain 
number are poisoned by the natives every winter. 

According to Sowerby (1923, p. 31) , this Tiger occurs throughout 
the forested areas of the Amur and the Ussuri, into Primorsk in the 


extreme east. It is said to be most plentiful in the Amur Province, 
round the mouth of that river; it is also numerous in the Ussuri 
Valley. Westward it probably extends almost to the Yablonoi 

"North of Khabarovsk they are extremely rare in the East, though 
I understand there are a few in the Bureya Mountains. We saw 
the tracks of only one in the region of Troitskov. In the Ussuri 
River region they seemed to be relatively plentiful. East of Bikin 
the forest seemed to be well tracked with tiger trails, but one tiger 
throughout the winter can make a lot of tracks. We secured three 
tigers fifty miles east of Bikin, during the winter 1929-30. As far 
as I could learn, these were all the tigers taken in this region during 
that winter. That the tigers have been able to hold their own up 
to now seems somewhat encouraging, though the present extended 
lumber activities of the Soviet Government take many Russian 
hunters into the forest. Previously about the only people who 
hunted the tiger were the Tungus tribes, with their primitive traps 
and snares. I, personally, am under the impression that it is only 
a matter of time until the tigers are reduced to the point of ex- 

"Tigers bring a big price in China, as medicine, but the hunter 
has to cover a lot of territory and work hard to get even one animal." 
(G. G. Goodwin, in litt., May 18, 1937.) 

Referring to the Maritime Province, Sowerby says (1934c, p. 40) : 
"Tigers of the long-haired species, whose skins are so valuable, were 
being secured in greater numbers than before, for, whereas formerly 
about ten of these great cats were killed in the province each year, 
over twenty had been killed during the first three months of the 
present year." 

W. G. Heptner writes (in litt., December, 1936) that more than 
ten are killed each year in eastern Siberia. Hunting is allowed 
throughout the year. Protection is given, however, in the reserve of 
Sikhota Alin. 

"Schrenck (1859, pp. 95-96) reported Fells tigris from Sakhalin 
as a rare winter visitor from continent, but his statement seems very 
doubtful" (Kuroda,1928,p.226). Ford Barclay (1915, p. 225) could 
find no evidence of its occurrence there. 

Manchuria. In this country, says Sowerby, the Tiger is "the most 
dreaded of the carnivores." He continues (1923, pp. 30-32) : 

His thick winter coat fetches a high price in the fur-markets of the world, 
being worth far more than those of the Bengal, Persian or Sumatran tigers. 
Not only is his skin of value, but his whole carcass; for the Chinese believe 
that the bones, blood, heart, and even the flesh of the tiger have medicinal 
properties of rare power, and will pay a goodly price for decoctions brewed 
by the apothecary that contain such ingredients as powdered tiger's knee-cap, 


or clotted tiger's blood. The heart of the tiger is supposed to impart to the 
consumer the courage and strength of the tiger itself. 

On this account the tiger has been hunted till he is almost extinct in 
most districts of North China, where once he was common, and now 
survives, even in Manchuria where he was once plentiful, only in the more 
remote and inaccessible forest areas, such as the Ch'ang-pai Shan, the Khingan 
Mountains, or the more or less unexplored and thinly settled areas of the Amur 
and Ussuri. . . . 

Formerly the tiger was extremely plentiful in all the forested areas of 
Manchuria. Indeed, it is said, they were so plentiful along the route of the 
western portion of the Chinese Eastern Railway when under construction, 
that they became a positive pest, killing and carrying off workmen, till a 
regiment of Cossacks had to be sent to cope with the situation. . . . 

In the forests of North Kirin and in Central and Western Heilungkiang 
tigers are killed by the local hunters every winter. 

The same author adds (p. 33) that the Russian hunters in Man- 
churia track the Tiger down in the snow, camping on its trail and 
following it for as much as ten days or a fortnight. The Chinese 
usually employ traps, pitfalls, and poison. 

Mongolia. In view of the fact that one of the names applied to 
the present subspecies is "Mongolian Tiger," it is surprising to find 
such a dearth of definite records from that wide country. According 
to Ognev (1935, p. 292), Radde (1862) reported the Tiger from the 
district of Uriankhai (the present Tannu-Tuva) , but later explorers 
have not found it there. (The subspecies of this region has not been 
determined.) Various other references in the literature to Tigers 
in Outer Mongolia give no information as to specific localities. 
Apparently the only likely areas for their occurrence are in north- 
eastern and eastern Mongolia, along the Siberian and Manchurian 

Korean Tiger; North China Tiger 


Felis tigris coreensis Brass, Nutzbare Tiere Ostasiens, pp. 4-5, 1904. (Korea.) 

(Fide Kuroda, 1938, p. 40.) 
FIGS.: Ford Barclay, 1915, pis. 84, 85; Sowerby, 1923, pi. 2 (coreensis^); 

Sowerby, 1933, pi. facing p. 166; Ognev, 1935, pp. 285-286, figs. 129-131. 

This Tiger apparently occurs in small numbers from Korea and 
southern Manchuria westward through the eastern border of Inner 
Mongolia and through North China. Its southern limits, where it 
presumably intergrades with the South China form, are not definitely 
known but perhaps may be roughly fixed at the divide between the 
Hwang Ho and the Yangtze Kiang Basins. 

The North China form differs from the Manchurian Tiger "in 
being smaller,- much darker and more fully striped and in having a 
shorter less woolly winter coat" (Pocock, 1929, p. 531). 


Korea. "In the Korea great value is apparently placed upon the 
skins, which are reserved for the chiefs" (Elliot, 1883, text to pi. 3). 
Ford Barclay (1915, pp. 228-231) gives the following account: 

Tiger are probably more numerous in the north than in the southern part 
of Korea .... 

In the neighbourhood of the foreign mining concessions, near the Yalu, 
dynamite is or was used with some success by native hunters, a small, specially 
constructed bomb being somehow concealed in the bait. Lately, however. 
. . . the Japanese police have forbidden the supply of dynamite for this 
purpose. Drop traps, weighted with stones and huge logs, are very common, 
and many tigers are accounted for in this way every year. 

[In Manchuria] the natives lay down poison wholesale. This is forbidden 
now in Korea .... 

My own most successful hunts have been in the island of Chindo, . . . 
situated at the south-west corner of Korea. . . . Early this year (1914) the 
body of a tiger was washed up on the west coast of Japan south of Matsue, 
at least 120 miles from the nearest mainland, from whence alone it could have 
come; yet, as reported in the press, its condition was such that the skin was 
removed for dressing and parts of the flesh sold for consumption 1 . . . 

This demand for tiger flesh on the part of the Japanese is a curious survival 
of barbaric superstition in such a highly civilized race. One of their chief 
officials sent me an urgent request for a shoulder on hearing of a successful 
hunt. This joint for some reason is supposed to possess greater medicinal 
virtue than any other, and the shoulder blade ground to powder is a certain 
cure in the most advanced stages of insanity! 

When a tiger is killed [in Korea] notice is at once sent to the elders of all 
villages within a radius of five miles, [and on their arrival a] wrangle ensues 
as to who are to be the privileged half-dozen to partake of a cupful of the 
ambrosial liquid left in the abdominal cavity, after the removal of the 
intestines. . . . 

Among both Chinese and Koreans, tiger's blood is believed to have an 
extraordinarily rejuvenating effect, greater even than the highly prized wapiti 
or sika horn .... 

Of the twenty odd skins I have seen in South Korea all have been much 
darker in colour than the half-dozen brought for my inspection in East 
Siberia .... 

In the happy days before the Japanese occupation and the consequent 
confiscation of fire-arms, when the depredations of a tiger became too pro- 
nounced, the active male inhabitants of the villages in the neighbourhood, 
perhaps half a dozen, armed with matchlocks, and as many more with heavy 
spears, would arrange for a day or two's driving in the adjacent hills. Occa- 
sionally these hunts were successful. 

"In North Corea tigers are said to be still fairly numerous, and 
every year some are killed there by sportsmen" (Sowerby, 1923, 
p. 31). 

In 1922 Kermit Roosevelt (in Roosevelt, Roosevelt, Derby, and 
Roosevelt, 1927, pp. 41-84) undertook an extensive but unsuccessful 
Tiger hunt with beaters in various localities of northern Korea. Some 
old tracks were found, but apparently the species is by no means 
common there. 

Manchuria. Sowerby 's records (1923, p. 30, pi. 2) from the 


Ch'ang-pai Shan, close to the Korean border, may refer to the 
present subspecies rather than to longipilis. 

Inner Mongolia. "A stuffed tiger's skin used to repose in a temple 
in Lama Miao (Dolonor) .... It was said that the animal . . . 
was killed in the streets of Lama Miao itself, having wandered from 
the Wei-ch'ang, or Hunting Grounds, to the east of that town." 
(Sowerby, 1923, p. 32.) 

China. Sowerby (1923, pp. 31-32) writes of this Tiger in China: 

How far west it extends is difficult to say, but it certainly reaches the 
western border of the province of Shansi, in North China, and southward 
reaches at least to the middle of the southern half of that province. From 
there it extends northward into Mongolia and in a north-easterly direction 
through Chihli, where it still occurs in the wilder parts of the Tung Ling 
and Wei Ch'ang (the Eastern Tombs, and Imperial Hunting Grounds) to 
the North-east and North of Peking .... 

In North China the tiger is becoming increasingly rare. In 1909 I saw the 
tracks in the snow of what must have been a very large animal in the 
mountains of West Shansi, in the Ning-wu district. I also heard of tigers 
in the Ko-lan Chou area and the Chao-ch'eng Shan, both heavily forested 
districts further south in the same province. Further south still near P'ing-yang 
Fu a tiger was killed by the natives about the year 1912. I have seen skins 
of tigers that were said to have come from the Kuei-hua Ch'eng area in North 
Shansi, and they were undoubtedly of the true long-haired type. The natives 
in this area also insisted that tigers occurred there. . . . 

According to Chinese accounts tigers also exist in Kansu, and on the 
Thibetan border, but I have been unable to get any satisfactory verification 
of this. It is more than likely that these animals occur for a considerable 
distance west of Kuei-hua Ch'eng into that little known mountainous country 
leading to the Ali Shan. 

In the early part of the present century an old native hunter 
reported the occurrence of three Tigers in the Eastern Tombs forest, 
in Hopei, during his lifetime. In 1932 a Tiger was killed after it 
had invaded a shop in the Yu Hsiang district of South Shansi. 
(Sowerby, 1933, pp. 167-168.) 

Owing to lack of specimens, it has not been determined whether 
the occasional Tigers reported in Szechwan (cf. Wilson, 1913, 
pp. 178-179, and Weigold, 1924, p. 74) belong to the North China 
or to the South China form. ^ 

"Tiger-bones ... are a highly prized Chinese medicine, and are 
supposed to transmit vitality, strength, and valour to those who 
partake of them. In the Imperial Maritime Customs Trade Returns 
of Hankow for 1910 is the following item: 'Tiger-bones, 77 piculs; 
value, Tls. 6522.' " (Wilson, 1913, p. 179.) 

"It is problematical whether or not predatory animals should 
be protected in a thickly populated country like China, but it seems 
a pity that such fine carnivores as the Chinese tiger (Panthera tigris 
styani Pocock), the Amoy tiger (Panthera tigris amoyensis Hilz- 


heimer) [,] the Manchurian tiger (Panther a tigris amurensis, Dode) 
. . . should not be preserved as part of this country's wonderfully 
rich mammalian fauna" (Sowerby, 1937, p. 257). 

[In South China, from Chekiang and Hupeh southward, and also 
westward to Yunnan, the Tiger occurs somewhat more commonly 
than in the more northerly regions. To this form Hilzheimer has 
given the name of Felis tigris var. amoyensis (Zool. Anz., vol. 28, 
p. 598, 1905; type locality, presumably the vicinity of Hankow, 
Hupeh) . It is recognized by G. M. Allen (1938, p. 480) , who regards 
Panthera tigris styani Pocock (1929) as a synonym. W. L. Smith 
(1920, pp. 355-363) gives an extremely interesting account of the 
methods of the native hunters in the vicinity of Amoy, who, armed 
only with torches and trident spears, track the Tigers into caves. 
There is also an account of the Tiger of Fukien by Andrews (in 
Andrews and Andrews, 1919, pp. 44-66) . 

In French Indo-China, Siam, and the Malay Peninsula the Tiger 
seems to be moderately common. For example, the number in Cochin 
China is estimated at 200-300; here it is of interest from the point 
of view of big-game hunting, but not commercially (Roche, Chef 
du Service Veterinaire du Cochinchine, in litt., 1937) . Rodolphe M. 
de Schauensee informs me that the Tiger is common in Siam (Thai- 
land) but preys chiefly on the wild game and does not seem to be 
regarded as a serious pest. The Tiger of these regions is not dis- 
tinguished by Pocock (1929, pp. 532-533) from the Indian Tiger. 

The Indian or Bengal Tiger (Panthera tigris tigris (Linnaeus)) 
ranges westward through Burma to India, where it inhabits the 
greater part of the Peninsula from the southern slopes of the 
Himalayas southward, but avoids the treeless and desert areas. It 
is not yet rare enough to call for any special discussion in this 

Tiger of Chinese Turkestan 


Felis tigris lecoqi Schwarz, Zool. Anz., vol. 47, no. 12, p. 351, 1916. ("Gebiet 
von Kurla, Lop-nor-Gebiet" (probably near Bagrash Kul), Chinese 

All the Tigers of Chinese Turkestan will be treated for convenience 
under this name, although the exact limits of the subspecies are 
unknown. Evidently the animal is not at all numerous, and its 
numbers may have declined to the point of extinction. In his review 
of the Tigers, Pocock (1929) seems to have overlooked the name 
of this subspecies as well as the occurrence of any Tiger in Chinese 


This is a very brightly colored Tiger, with regular pattern, fore- 
legs unstriped in front, conspicuous shoulder tufts, short neck mane, 
thick cheek whiskers, slightly lengthened abdominal hair, and a thick 
winter pelage. It differs from the form of Russian Turkestan in 
having smaller stripes and dull brown instead of black thigh mark- 
ings. (Schwarz, 1916, p. 352.) 

J. H. Miller (in Carruthers, 1913, pp. 582, 609-610) writes of this 

The dense jungles which cover so large a portion of the [Dzungarian] 
lowlands . . . are the haunts of the tiger .... 

The tiger inhabits the same country as the wapiti, though, perhaps, keeping 
rather more to the dense reed-jungle. It is, however, not entirely restricted 
to the plains, for in the Kash, Kunguz, and Jingalong valleys, on the Upper 
Hi River, it is found at an altitude of from 4,000 to 5,000 ft. among the 
thick scrub on the edge of the spruce forest. Every year a few tiger-skins 
find their way into the Urumchi, Manas, or Shi-Kho bazaars. They are, in 
nearly every case, secured in winter, by the farmers and herdsmen living 
on the edge of the jungle, by means of poisoned carcasses of sheep or goats. 
Very few of the natives would dare to fire at a tiger .... Wild-pig . . . are 
undoubtedly the tigers' staple food, but during the winter they occasionally 
raid a farmer's flocks, and it is then that poisoned carcasses are laid out for 
them. . . . 

I doubt if they are anywhere numerous. . . . 

It must be remembered that the tiger which inhabits Dzungaria and the 
Tarim basin, also the Ala Kul, Balkash, Syr Darya, and other portions of 
Russian Turkestan, is a very different animal to the Manchurian variety. 
It is not so long-haired, and it is considerably smaller and less finely marked. 

Theodore Roosevelt (in Roosevelt and Roosevelt, 1926, p. 166) 
writes of Tigers in the Tian Shan: "We were told that they existed 
no longer in the Tekkes. . . . They [the natives] said that during 
the last ten or fifteen years the native hunters had killed them off 
with poisoned meat." 

"The tiger . . . formerly ranged in the forests on the edges of the 
Tarim Basin and the swampy areas along the northern slopes of the 
Thian Shan. . . . The tiger seems to have been exterminated.'' 
(Morden, 1927, p. 123.) 

Alpheraky (1891) reported the species from the Tekes and the 
lower Kunges, tributaries of the Hi River in Dzungaria (Ognev, 
1935, p. 291). 

"The . . . tiger, which formerly inhabited the woods of the middle 
Tarim, seems to be dying out" (Hedin, 1940, p. 149) . 

Caspian Tiger; Persian Tiger 


Felis virgata Illiger, Abhandl. K. Akad. Wissen. Berlin, 1804-11, physikal. 
KL, pp. 90 and 98, 1815. ("In Persien und am Kaspischen Meere"; type 
locality restricted by Harper (1940, p. 194) to the "Province of Mazanderan, 
northern Persia.") 


SYNONYMS: Felis (Tigris} tigris septentrionalis Satunin (1904); Felix tigris 

trabata Schwarz, 1916. 
FIGS.: Heck, Lebende Bilder, p. 157, 1899; Kennion, 1911, pi. facing p. 251; 

Pocock, 1929, pi. D, lower fig., pi. 3; Ognev, 1935, figs. 121-124. 

While the Indian or Bengal Tiger (Panthera tigris tigris x ) prob- 
ably exacts a greater annual toll in human lives than any other 
carnivorous mammal, some of its races in western, central, and 
eastern Asia and in the Malay Archipelago conflict to a far less 
degree with the interests of mankind. In any event, their numbers 
have been reduced to a point where they are entitled to a place 
in the present work. 

The Caspian race is "generally a medium sized or smallish tiger 
with a thick longish winter coat, dark in colour, with numerous, 
close-set stripes showing a marked tendency to brownness on the 
whole or parts of the body." Length of male, about 10 feet 8 inches; 
of female, about 8 feet 6 inches. (Pocock, 1929, pp. 522, 540.) 

The range of this Tiger extends from Transcaucasia (formerly) 
through northern Persia to northern Afghanistan; presumably the 
same form occurs northward to the Aral Sea and Lake Balkash in 
Russian Turkestan (formerly to the upper Ob Basin and the Altai 
region) . 

Transcaucasia. "A few are annually killed in Turkish Georgia" 
(Blyth, 1863, p. 182). 

Satunin reports (1906, pp. 308-309) as follows on the Tigers of 

At the time of Radde's first expedition to Lenkoran in 1866 Tigers 
were still very numerous there. In seven weeks six fresh skins were 
offered him. But in 1879-80, in the course of eight months, he 
could not secure a single fresh skin. Tigers still occurred, but were 
very rare. According to the hunters' reports, the animals were quite 
extirpated somewhat later, but in the 90's they began to increase, 
and at the time of my expedition (1897-99) two to four specimens 
were taken annually. At present they occur chiefly in the Prisib 
district of Lenkoran, both in the lowland forests and in the foot- 
hills. In 1899 Tiger tracks were found on the Mugan Steppe, where 
the animal had gone apparently in pursuit of Wild Boars. 

Satunin also expresses here the conviction that the numerous 
reports of Tigers in other localities of Transcaucasia are due to 
a confusion of this species with the Leopard. In a previous paper 
(1896, pp. 289-290) he had stated that they occurred formerly as 
far as the ridge of the Great Caucasus, and he had quoted Nord- 
mann's report of Tigers killed near Tiflis in 1835. The species is 
now exterminated in Transcaucasia (W. G. Heptner, in litt., Decem- 
ber, 1936). 

i Fells tigris Linnaeus, Syst. Nat., ed. 10, vol. 1, p. 41, 1758. (Bengal.) 


Persia. Gmelin (1774, vol. 3, pp. 485-486) reports the animal 
as pretty common in the forested mountains of Mazanderan. It 
seldom makes unprovoked attacks on man. The skin is highly 
prized, and is used for a horse-covering. 

Blanford writes (1876, p. 34): "The tiger is only found in 
Persia in the Caspian provinces, Mazandaran, and Ghilan, lying to 
the north of the Elburz mountains .... These provinces, unlike 
the plateau of Persia, are covered with dense forest, and in them 
the tiger ranges up to an elevation of at least 5000 or 6000 feet." 

To this St. John adds (in Blanford, 1876, p. 34) : "Tigers are 
very numerous in the Caspian provinces of Persia, and in the 
Caucasus as far as the mouth of the Araxes. . . . Cubs are often 
captured in Mazandaran and brought to Tehran. I have seen speci- 
mens in the Bagh-i-Washi quite equal in size to Bengal tigers." 

In Mazanderan, sometime prior to 1911, "Col. Kennion only came 
across two examples of this tiger; and there is reason to fear that 
the race is on the wane" (Pocock, 1929, p. 522). "Considering the 
abundance of game and the fewness of the tigers' foes, it is quite a 
problem why the latter are not more numerous in these parts" 
(Kennion, 1911, p. 246). 

The British Museum has a specimen obtained at Astrabad in 1882 
or earlier. In Astrabad and the adjacent portion of Turkestan the 
Tiger occurs in various localities, including the Gurgan, Atrek, Sum- 
bar, and Chandir Rivers (Ognev, 1935, pp. 289-290). 

Afghanistan. In this country, as in Persia, the species appears 
to be restricted to the northern part. "The tigers of the Perso- 
Turkestan district . . . were doubtless excluded from India by the 
Hindu Koosh and the desert areas of Persia and Baluchistan" 
(Pocock, 1929, p. 509). 

"Ferrier in his 'Caravan journeys' speaks of tigers in the jungles 
of the Hari Rud north-west of Herat" (St. John, in Blanford, 1876, 
p. 34). 

The Afghan Delimitation Commission (1884-85) obtained a speci- 
men from Karaol-khana on the Murgab close to the Turkestan 
boundary. Tracks were reported in the valley of the Hari Rud, 
and were also found at the Chashma-sabz Pass, at an elevation 
of 5,000 feet, in the Paropamisus Range. "During summer . . . 
they wander over the great rolling plains of the Badghis [on the 
north side of the Paropamisus Range], ascending to higher altitudes 
with the increase of heat, depending for their food on Pig, Oorial, 
and even Ibex. In winter they resort to the . . . thickets of the 
larger streams and main rivers, to which their usual food, the Pig, 
also retires. The Turkomans say that an old and toothless Tiger is 
especially destructive to sheep." (Aitchison, 1889, p. 56.) 


Russian Turkestan and Western Siberia. Ehrenberg reports 
(1831, p. 389) that Tigers are frequently observed on the Tarbagatai 
Mountains southwest of Zaisan Nor; also that the Cossacks of the 
Irtish have several times killed Tigers on the Kirghiz Steppe, spear- 
ing them from horseback. 

"North of the Hindu Kosh, Tigers occur in Bokhara, and proved 
troublesome to the Russian Surveying Expedition on the shores of 
the Aral in midwinter. They are also found on the banks of the 
Irtisch, and in the Altai region." (Blyth, 1863, p. 182.) 

Atkinson (1858, p. 282) mentions four specimens in the museum 
at Barnaul, western Siberia. "The tigers were killed in Siberia at 
different places, some at a distance of about five hundred versts 
from Barnaoul ; they had come from the Kirghis Steppe, and crossed 
the Irtisch into the Altai in the region around Bouchtarminsk. . . . 
They are rarely found in Siberia; it is only when they are driven 
from the steppe by hunger that they cross the Irtisch most prob- 
ably when following the track of their prey : many peasants do not 
even know them by name." Atkinson also reports (p. 486) many 
Tigers about the western end of the Ala Tau, southeast of Lake 

According to Severtzoff (1876, p. 49), the Tiger "is common in 
Turkestan, especially up to about 4000 feet altitude; but beyond 
that it is rare in winter, and only in the summer does it visit localities 
which are higher than 7000 feet." 

Carruthers writes (1915, pp. 149-150) : "In the same locality 
[Oxus or Amu Darya Valley] inhabited by the Bokharan stags, 
tigers are fairly numerous. These we know range the whole course 
of the Oxus from the Sea of Aral to the foot of the mountains near 
Kulab. They are seldom hunted or seen. I have good reason to 
believe they wander across the desert from the Oxus to the lower 
Zarafschan. The natives speak of them, and I am certain I heard 
one one night in the saxaul forests which surround the swamps 
where the river loses itself in the sands, and where large numbers of 
wild pig roam." 

The British Museum has a skull from the vicinity of Find j eh, 
on the Murghab (Pocock, 1929, p. 522). 

In Turkestan the Tiger reaches its northwestern limit at the 
Gulf of Karabugas on the Caspian Sea, avoiding the Ust Urt Plateau. 
It was formerly numerous on the Murghab and Tejend Rivers, the 
last having been killed in that region in 1904. During a period of 
some years prior to 1915 nine Tigers were killed in the Syr Darya 
region. The species also occurs in the valley of the Chu and on the 
Amu Darya delta. In 1887 it was reported as abundant on the lower 
Hi River and on the southeastern shore of Lake Balkash; by 1930 
its numbers in this region were few. There are old records from the 


Tarbagatai Mountains southeast of Zaisan Nor, and from Zmeino- 
gorsk, Bisk, and Barnaul in the Ob Basin of western Siberia. The 
Tiger has entirely disappeared from its former haunts in the Dzun- 
garian Alatau. In Tajikistan it occurs on the upper Vashni and on 
the Kafiringan Darya. (Ognev, 1935, pp. 273, 290-292; map, p. 295.) 
The following information is from W. G. Heptner (in litt., Decem- 
ber, 1936) : The Tiger is found in limited numbers, but regularly, 
at the mouths of the Amu Darya and the Syr Darya and on Lake 
Balkash. It is more common on the upper course of the Amu Darya 
and on its right tributaries. It comes over accidentally from Persia 
and Afghanistan to Kopet-Dag, the upper Tejend, the upper Mur- 
ghab, and Transcaucasia (Talish) . During the past 50-70 years the 
numbers have been considerably reduced by hunting. The range 
has also been reduced, and in certain areas (Transcaucasia, middle 
Syr Darya, and Murghab) the Tiger is now exterminated. It is 
difficult to estimate the total number, but there may not be more 
than 200 in Russian Turkestan. The best areas are the headwaters 
and the mouth of the Amu Darya. The Tiger is rarely met with 
at the mouth of the Hi River on Lake Balkash, where probably only 
ten or twelve animals exist. At the mouth of the Syr Darya it is 
probably only a visitor, coming from Amu Darya. Hunting is 
allowed throughout the year. 

Javaii Tiger , 


Tigris sondaica Fitzinger, 1 Sitz.-ber. Akad. Wiss. [Wien], math.-nat. Cl., vol. 68, 
pt. 1, p. 454, 1868. ("Java und Sumatra"; type locality restricted by 
Schwarz (1912, p. 324) to Java.) 

The meager information available concerning the status of the 
Javan Tiger indicates that it is no longer very numerous or generally 
distributed on that island. 

"Ground-colour light rusty; stripes very narrow, often duplicated. 
. . . Fur short and close." (Schwarz, 1912, p. 325.) "Apparently 
closely resembling the Sumatran race in size and coloration, but 
distinguished from it, and from all other tigers, by the marked con- 
striction of the occiput" (Pocock, 1929, p. 541). 

In 1851 Horsfield (p. 44) remarked that Tigers were "numerous 
and destructive ... in many parts of Java." 

"Many tigers . . . may be found" on the Oedjoeng koelon Penin- 

i This name is antedated by Felis tigris sondaicus Temminck (Coup-d'oeil 
Possessions Neerlandaises, vol. 2, p. 88, 1847). It is highly questionable, how- 
ever, whether Temminck's excessively brief and insufficient description ("le 
grand tigre raye de Sumatra et de Java forme une espece distincte du tigre raye 
du continent de FInde") is nomenclaturally valid. 


sula, at the extreme western end of Java, which constitutes a nature 
reserve (Dammerman, 1929, p. 34) . 

"In Java, the tigers living up to 1914 in the swamp country near 
Maoek, are now extirpated. In 1931 they were seen on the Goenoeng 
Malabar. They are also found in the Baloeran District, southern 
Banjoewangi, in the Southern Mountains, and near Banjoemas. Two 
to four are shot every year at Tampomas. Finally a number of 
tigers are also reported from S. E. Garoet" and from the Midangan 
district. (Heynsius-Viruly and Van Heurn, 1936, p. 58.) 

[The Sumatran Tiger (Panthera tigris sumatrae *) , although less 
common than formerly, is still numerous in various districts, and its 
protection is not urged at present. (Heynsius-Viruly and Van Heurn, 
1936, p. 59).] 

Bali Tiger 


Felis tigris balica Schwarz, Ann. Mag. Nat. Hist,, ser. 8, vol. 10, p. 325, 1912. 

("Den Pasar, Siid-Bali.") 
FIG.: Schwarz, 1913, p. 71, fig. 

In Bali the Tiger seems to be in rather imminent danger of 

It is very similar to the Javan Tiger, but smaller; ground color 
somewhat brighter, and the light markings clearer white; fur short 
and close. Head and body, 1,530 mm.; tail, 580 mm. (Schwarz, 
1912, p. 326.) 

About 1909-12 the Tiger was considered fairly common in Bali; 
yet information concerning damage done by it was not forthcoming 
(Schwarz, 1913, p. 73). 

"A few yet live in West Bali, but they are having a hard time 
because they are much sought by hunters from Java, so that they 
will certainly disappear within a few years. The species also exists 
in N. W. and S. W. Bali." (Heynsius-Viruly and Van Heurn, 1936, 
p. 58.) 

Order PROBOSCIDEA: Proboscideans 

Family ELEPH ANTID AE : Elephants 

The Elephants are composed of an Asiatic genus (Elephas) and 
an African genus (Loxodonta) . Lydekker (1916) recognizes 4 
Asiatic forms and 11 African forms, but Dr. Allen (1939b) ques- 

i Panthera tigris sumatrae Pocock, Jour. Bombay Nat. Hist. Soc., vol. 33. 
no. 3, p. 535, pi. H, 1929. ("Deli, Sumatra.") 


tions the taxonomic status of all but 4 in the latter group. The 
distribution of the family covers southeastern Asia (India, Burma, 
Ceylon, Thailand, French Indo-China, Malay Peninsula) , Sumatra, 
Borneo (introduced?), and the greater part of Africa south of the 
Sahara. Accounts of three forms are supplied herein. 

Malay Elephant 


Elephas maximus hirsutus Lydekker, Abstr. Proc. Zool. Soc. London, no. 130, 
p. 20, 1914. ("Kuala Pila district of the Negri Sembilan province of the 
Malay Peninsula" (Lydekker, 19146, pp. 285-286).) 

FIGS.: Lydekker, 19146, p. 285, fig. 1; Lydekker, 1916, vol. 5, p. 84, fig. 25. 

The Elephant of the Malay Peninsula is regarded by competent 
authorities (e. g., F. N. Chasen, in litt., March 31, 1937) as "a 
vanishing form." 

This subspecies is "characterized by the square, instead of tri- 
angular, form of the ear, the early date at which its upper margin 
is bent over, and the presence in the young condition ... of a 
thick coat of black and in part bristly hair" (Lydekker, 1914a, 
p. 20). 

The northward range of the Malay Elephant has not been de- 
termined; it will here be provisionally considered to extend as far 
as the Isthmus of Kra, in Peninsular Siam. In the remainder of 
Siam and in French Indo-China the Elephant belongs presumably 
to the Indian subspecies and is reported as more or less common 
(Gyldenstolpe, 1919, p. 169; James L. Clark, in litt., June 26, 1936; 
P. Vitry, in litt., December, 1936; Roche, in litt., 1937). 

Malay States Flower says (1900, p. 365) : "Wild elephants do 
not occur in either Penang or Singapore, nor are tame ones em- 
ployed there; but on the continent, both in Siam and the Malay 
Peninsula, elephants are found wild in suitable localities, and are 
trained for various purposes. ... I saw more or less trained 
elephants in ... Kedah, and Perak, but in the Southern Malay 
States the people do not seem to catch and tame them." He also 
(p. 366) quotes H. J. Kelsall (1894) to the effect that "the elephant 
appears to be common throughout Johore"; and H. N. Ridley 
(1894) as remarking that "the elephant, though common all through 
Pahang, is never caught and tamed." 

Referring to conditions from 1900 on, Burgess writes (1935, 
p. 249) : "Elephants roam all over the peninsula and are common as 
far south as Johore. . . . Since only a small fraction of the jungle 
has yet been cleared, the probabilities are that large herds have 
not been seen." 


Hubback (1923, pp. 24-25) reports on "the damage done to plan- 
tations by elephants" in the Malay Peninsula: 

It is a very extraordinary thing, but all wild animals which browse seem 
to acquire an unholy craving for the bark and leaves of Hevea brasiliensis. 
Wild elephants especially, once they have tasted the bark, seem to go mad 
for it. I have absolutely trustworthy evidence of an eye-witness who has 
seen elephants strip the bark, from rubber-trees by first catching hold of a 
small piece with the tip of the trunk and then pulling upwards, so that 
a strip of bark is taken off the tree. In an incredibly short time the tree is 
ruined. Then they love to lean against the trees, and I suppose are sur- 
prised and annoyed when they fall over. Undoubtedly these wild elephants 
are in certain places a serious nuisance. 

A further account is given by Hubback in the Report of the Wild 
Life Commission of Malaya (vol. 2, 1932) . He says that elephants 
are not uncommon south of Gunong Sinting, between that mountain 
and the Pahang border, and continues: 

It is a fact beyond question that wild elephants do and have done consider- 
able damage amounting to values of thousands of pounds. Had it not been 
for elephants in Malaya still larger areas planted with rubber would now be 
yielding latex. These facts are not in dispute. . . . 

The elephants known as the "Carey Island Herd," which lived on a large 
island on the coast of Selangor, which island was given out for agriculture, 
were all ultimately destroyed. Their death warrant was really signed when 
the grant for the land was made out. Then there is the "Kuala Selangor 
Herd" which has been almost totally exterminated; a cow and a calf being 
reported as the sole survivors. The destruction of the survivors was advo- 
cated. This herd must have consisted of 40 or 50 animals thirty years ago. 
The "Labu Herd" in Negri Sembilan has been practically eliminated. In 
Lower Perak the "Chikus Herd" of elephants has given a lot of trouble and 
many of them have been shot. In many other places elephants have been 
harried and driven from locality to locality in alleged defense of agriculture. 
The records of elephants that have been killed in Malaya during the last 
few years under the agriculturist's exemption are incomplete reports are 
seldom sent in of elephants that have been wounded but there is reason to 
believe that the Malayan elephant is on the way to extermination. It is 
extremely doubtful if the yearly toll of destruction is being made up by the 
yearly production of calves, and that means extinction unless a halt is called. 
One must take into account the fact that wild animals when much disturbed 
have a habit of curtailing their breeding, and it is almost certain that this 
affects elephants as well as the other large forms of our fauna. 

In Kuala Selangor, Lower Perak, Labu, and elsewhere, despite the sup- 
posed sanctuary provided by Forest Reserves, the elephants have not been 
left undisturbed, and have been unable to find in the areas that they have 
receded to that tranquility essential to an elephant's well-being. 

Where elephants have been forced to live in jungle areas which are insuffi- 
cient for their normal existence, and where they have become a serious 
menace to cultivation, it is advocated that they should be destroyed by per- 
sons whose business it would be to undertake the work. . . . 

It is an established fact that wild elephants, always providing they are not 
suffering from wounds, can be driven away by fire crackers and noise. In 
cases of absenteeism, which is frequently the contributory cause when ele- 
phants visit native cultivation, these methods cannot be applied. A woven 


wire fence properly upkept and with a path kept reasonably clean on the 
jungle boundary . . . would in most cases keep elephants from entering the 
cultivated area. . . . 

The removal of protection from Elephants, a measure taken in 1929, was 
condemned by the vast majority of English speaking witnesses before the 
Wild Life Commission of Malaya. This order was liable to accentuate the 
trouble from wounded elephants and undoubtedly, as evidence showed, forced 
elephants into localities where they had never been known before. This 
unwise order was rescinded on the 15th of May, 1931, and the elephant 
cannot now be shot at by an unlicensed person except in alleged defence of 
property. . . . 

Raids on native cultivation are often due to neglect. Persons familiar with 
the habits of elephants can often move a herd from the vicinity of cultivation 
by following them up all day until they are miles away from the locality 
they visited the previous night. . . . 

How do Sakai in their primitive state handle the planting of crops in ele- 
phant country? Showing more wisdom than their white brothers, they leave 
elephants alone. ... In the Sakai country, which lies between the main 
range and the Kelantan Railway, the Sakai suffer no damage from elephants. 
The elephants, not being disturbed and harried, have not learned to "answer 

[Some hold an opinion] that a very large percentage of the so-called damage 
done by elephants is only done to patches of abandoned cultivation, and 
when inhabited land is attacked it is not infrequently done by bad-tempered 
elephants suffering from wounds of sorts which are caused by some home- 
made bullets fired from a shot gun. 

F. N. Chasen writes (in litt., May 5, 1937) : "The question of 
protecting the elephant in the Malay Peninsula raises and crystal- 
lizes the whole policy of local big-game preservation. Can big-game 
co-exist with modern agriculturalists? My view is that the elephant 
should be protected in reserves : outside the reserves he must behave 
himself, or be shot. These are, of course, the extremes of the case 
and a middle course is, sometimes, permissible when directed by an 
experienced game-warden. The Malayan elephant is decreasing in 
numbers, rapidly, in the settled areas. It is still numerous elsewhere." 

Peninsular Siam. The following two accounts relate to the 
uninhabited country about the northern end of the Inland Sea : 

"On the plain and in the forest a herd of about 300 wild elephants 
are roaming. . . . These elephants have from time to time been 
captured, but their death has always resulted after some compara- 
tively short time." (Havmoller, 1926, p. 365.) 

"From government officials with whom I was traveling I learned 
that a herd of at least 200 elephants ranges over the vast grassy 
plain extending southward from near Nakon Sritamarat almost to 
Singora on the west side of the Inland Sea and practically from the 
Gulf of Siam to the high mountains in the west. This plain, suitable 
for rice growing, is entirely uncultivated owing to the ravages of the 
elephants." (H. M. Smith, 1926, pp. 365-366.) 

Elephants are protected in Siam because "they are considered 


property of the State, and therefore a special permit must be ob- 
tained from the King before an elephant may be killed" (David E. 
Kaufman, in litt., March 8, 1933). 

Suiiiatrun Elephant 


Elephas Sumatranus Temminck, Coup-d'oeil Possessions Neerlandaises, vol. 2, 

p. 91, 1847. ("Sumatra.") 
FIGS.: Lydekker, 1916, vol. 5, p. 83, fig. 24; Pieters, 1932, p. 58, fig. 

This Elephant, while still existing in considerable numbers in 
Sumatra, is evidently losing ground in contact with cultivation, and 
concern is felt over its future. 

It is said to be characterized by its small size, its tessellated 
skin, the pyriform shape of its ear, and the infolding of the posterior 
edge of the ear (Lydekker, 1916, vol. 5, p. 84) . 

Sumatra is the only part of the Malay Archipelago that has pos- 
sessed a native stock of Elephants within historic times. Those now 
found in Borneo are considered descendants of domesticated indi- 
viduals introduced from the Malay Peninsula (Mjoberg, 1930, 
pp. 15-16). 

In 1906 W. L. Abbott (in Lyon, 1908, p. 622) saw many trails in 
eastern Sumatra opposite Pulo Rupat. 

Only mature males may be hunted, and the open season may not 
exceed six successive months. The export of either living specimens 
or the skins of Elephants is prohibited, and the export of ivory is 
restricted within certain limits. During the past ten years an 
average of only 350 kilograms of ivory has been exported annually 
from the Netherlands Indies. The published value is only 10 to 20 
guilders a kilo. Animals with very large tusks have disappeared 
for the most part, and the present average weight of a pair of 
tusks is estimated at 10 to 12 kilograms. Thus the above-mentioned 
export figures represent the annual taking of about 35 Elephants. 
(Dammerman, 1929, pp. 13-14.) 

"The two principal ports to which the ivory is sent, are Singapore 
and Penang. Much ivory is also carved here locally, so we may 
suppose that yearly many more elephants are killed than the 35 
the tusks of which are exported. With the new regulations export 
of elephant-tusks weighing less than 5 kilograms a piece, is for- 
bidden." (Dammerman, 1929, p. 14.) 

The same author (in Skottsberg, 1934, p. 422) considers the 
Sumatran Elephant threatened with extermination. According to 
Pieters (1932, p. 58), the greatest danger is the encroachment of 
cultivation on its habitats. 

"There are still some elephants in Langkat District, but not as 


many as, say 10 years ago. There are some on Lepan, Besitang, 
and Namoe Oengas. Elephants are shifting from one place to an- 
other, and then come back to the starting point again." (J. Gourin. 
in Hit., August 7, 1933.) 

The following account is given by Heynsius-Viruly and Van 
Heurn (1936, pp. 48-50) : 

Very detailed reports on the elephant were received from many districts. 
While some believe they will be exterminated within the next twenty-five 
years, others think that they are holding their own as there are yet about 
2000 elephants in Southern Sumatra alone. This estimate is, however, called 
in doubt by competent observers. In the subdivision Ogan Oeloe there were at 
the most about 45 elephants in 1926. There is much difference of opinion 
about the damage these animals do. In Rokan they have increased so rapidly 
that they have become a nuisance; nevertheless they are not hunted much. 

A report from Soengi Radja relates that in 1929 a herd of 14 head was 
discovered; efforts are being made to preserve them. Elephants were also 
seen near Soengi Roka in May 1932. In Siak their number is estimated as 
still quite large, likewise in Indragiri, although they do not appear there 
in the swampy coastal districts. In the lowlands they are found only in 
Reteh, and the largest herds in South Seberida in the Boekit Tiga Pdeloeh. 
In the first-named district a reward of twenty-five Dutch guilders is offered 
for every elephant tail. The controler of the district, which comprises the 
middle course of the Siak River, paid twenty-eight such premiums in 1930. 
A correspondent estimates that about 200 elephants roam over Siak and 
urges the repeal of the old local regulation concerning the premiums, as 
well as not extending the permits for the fire-arms kept in the kampongs. 

In Djambi, Moeara Tambesi and Moeara Tebo they are fairly common; 
also even now, in South west Bangko, where the controler estimates they 
will be extinct within 10 years. They are very rare in Moeara Boengo. In 
Djambi they are estimated at about 250-350. 

There is a herd of about 30 in Korintjih, and seven in the Ophir district 
(July 1932) viz: one young male, and six females. In 1915 this same group 
numbered still 18. In 1916 the herd of North Korintjih were hunted by men 
specially appointed for the purpose by the Demang of Korintjih and the 
Civil Authority of Air Hadji. Not even the females and the young were 
spared at that time. Along the Mesoedjih River elephants are caught in 
pitfalls by the Natives and the younger ones are sold in Palembang. The 
older ones are left to starve, in order to obtain the tusks to sell. It is generally 
thought that the present regulations merely postpone the extermination of 
the elephant. Only establishing extensive reserves might bring adequate 
protection. . . . 

The report of a herd of 14 in Soengi Radja is of much interest, for these 
animals occupy a rather small area that has been completely surrounded by 
cultivation for a quarter of a century, and though much hunted they have 
succeeded in holding their own. The establishing 1 of a reserve here was urged 
in 1929. In 1932, the Netherlands Committee for International Nature Pro- 
tection requested this from the Government of the Netherlands Indies, but 
as yet no actual steps have been taken. 

A second important fact, emphasized by our enquiries, is that at certain 
seasons elephants migrate periodically from the mountains to the lower 
coastal areas. This too had been exhaustively recorded in print. It was one 
of the strongest arguments used by the Netherlands Committee, when sub- 


mitting their proposal for requesting the Indian Government to include in 
the reserve certain lowland swampy areas. This has not been done at present ; 
but the Committee intends calling the Government's special attention to the 
new information received which further supports their claim. The protection 
of the elephants remains, moreover, a separate problem. Even after re- 
serves are established, and, of course, after the Decree on Hunting becomes 
effective in Sumatra, the careful listing of existing herds will be imperative. 
The continued gathering of data regarding each herd, in order to determine 
which way they travel, their increase or decrease, and what damage they do, 
will furnish the foundation for their protection and for the preservation 
of the remaining herds. . . . May complete cooperation by the Department 
of the Interior facilitate this task of the Netherlands Committee. 

South African Bush Elephant 


Elephas ajricanus Blumenbach, Handbuch der Naturg., ed. 5, p. 125, atlas, 
pi. 19, fig. C, 1797. (Selected as Orange River, South Africa.) 

SYNONYMS: Elephas capensis F. Cuvier, Tableau Elem. de PHist. Nat. des 
Anim., p. 149, 1798 (Orange River region, South Africa) ; Elephas ajri- 
canus toxotis Lydekker, Proc. Zool. Soc. London 1907, pp. 385, 388, Aug. 1, 
1907 (Addo Bush, South Africa) ; Loxodonta africana zukowskyi Strand, 
Arch, fur Naturg., vol. 90, sect. A, pt. 1, p. 68, footnote, July 1924 (Kaoko- 
veld, South-West Africa). 

FIGS.: (Of the Tanganyika animal) M. Maxwell, 1930, 11 plates from 

Because of its great size, its strength, its tusks of ivory, its 
remarkable trunk, or proboscis, and its intelligence, the African 
Elephant is one of the most interesting of mammals. At first con- 
fused by early naturalists with the Indian Elephant, it is, however, 
so different that the two are now placed in separate genera. The 
African Elephants (Loxodonta) differ in many points, such as the 
structure of the tip of the proboscis, with its two instead of single 
fingerlike tips, the huge ears extending back to cover the sides of the 
neck as far as the shoulder blade, the forehead, which is less globular 
than in the Indian species, the cheek teeth having fewer of the high 
enamel prisms which form their essential grinding structure, the six 
successive teeth with usually 3, 6, 7, 7, 8, and 10 prisms, respectively, 
against 4, 8, 12, 12, 16, 24 in the Indian Elephant ( W. L. Sclater) . 
The African Elephant is slightly the larger, but seldom exceeds 
11 feet in height at the shoulder, a distance not easily measured with 
accuracy even when the animal lies dead. The weight of the famous 
"Jumbo" was about 6.5 tons. In color the skin is slaty gray but may 
appear in life of different tints, according to the light, the dryness 
of the skin, and the amount of earth clinging to it if animals have 
been dusting or wallowing. A sparse coat of short stiff hairs is 
insufficient to obscure the hide, but near the tip of the tail these 
hairs become stout coarse bristles growing from the edges of the 


compressed terminal part, several inches in length. The upper pair 
of incisors are enormously enlarged to form tusks, which are larger 
in the male than in the female, or may in the latter sometimes be 
lacking. These are used as weapons or in digging for roots. The 
largest tusks come from Kenya Colony, with a record length on the 
outside curve of 11 feet 5^ inches, and a weight for the two of 
293 pounds (Roland Ward, 1935) . 

The African Elephants have at various times been subdivided into 
local races by systematists, but there is still much doubt as to the 
value of the characters claimed, and the number of valid geographical 
forms. In general one may distinguish the larger "Bush Elephants" 
and the smaller "Forest Elephants," the former distinguished by 
minor additional points such as the larger, more elongated ears, the 
more forwardly directed tusks, less abundant hair. There is a ques- 
tion whether these two types should be regarded as merely races or 
as separate species, but the likelihood is that they have evolved 
side by side though in different habitats, the former avoiding the 
denser forests, the latter keeping more strictly to their shelter, with 
the result that at present the two types seem different enough for 
separation as distinct species. The larger Bush Elephants, again, 
have been regarded as of several local races, of which that of South 
Africa, the first to be named, is at present much reduced in numbers. 
Farther to the northeast, the East African animal has been named 
L. a. knochenhaueri, and the Sudanese Elephant, L. a. oxyotis. There 
is still much doubt as to the validity of the characters distinguishing 
these races, but until series of skulls and measurements can be com- 
pared one can only await further information. The character of the 
ear lobe invoked, for example by Lydekker, is so subject to modi- 
fication through distortion in dried or mounted specimens that 
little reliance can be placed upon it. One may then consider the 
status of the Bush Elephants as a whole, with special reference to 
the South African race. 

In classical times elephants were found over most of Africa except 
the most desert areas. There seems to be evidence that in ancient 
times they were found abundantly in Abyssinia, for under the 
Ptolemies, in the third century B. C., elephants for use in warfare 
were captured and trained in Ethiopia on the shores of the Red Sea 
and were taken thence in specially constructed boats to Egypt. 
Entire army corps were sometimes engaged in their capture. In 
Carthaginian days elephants were captured in Libya and in Maure- 
tania among the forest-covered foothills of the Atlas Mountains. 
Here, however, they have long ceased to exist and are not now found 
north of the southern borders of the Sahara. In the eastern Sudan 
elephants still occur in small numbers (I myself saw their "sign" 
on the Blue Nile, near the Abyssinian border in 1913), but have 


long ago retreated from the borders of the Red Sea. Swayne wrote 
a quarter of a century or so ago: "There is practically no elephant 
shooting to be got in Somaliland north of the Haud Plateau, or in 
the Haud, at the present time. In the gorges which descend from 
the highlands of Abyssinia to Ogaden in the country about the 
head-waters of the Webbi Shabeyleh and Juba Rivers there are 
still plenty of elephant. A few herds, it is believed, wander down 
those river valleys to the Marehan Country far to the south-east 
of Berbera." A certain amount of ivory-hunting by natives may 
keep these herds in check. But recent travelers up the Nile report 
large numbers of elephants in the practically impenetrable papyrus 
swamps of "the Sudd" where they will doubtless find sanctuary for 
a long period to come. Between this area and Uganda there are 
large numbers of elephants, and in the Kenya forests and thorn- 
bush a good many still survive. 

For the purposes of the present report, chief interest centers in the 
elephant of South Africa, which nowadays with increasing settle- 
ment of this part of the continent comes into close association with 
white men and has had to suffer in consequence. The following brief 
notes are given in summary from Shortridge (1934, vol. 1, p. 362) and 
W. L. Sclater (1900) . This, the typical race of African Elephant, at 
present seems to be characterized in part by its rather short stout 
tusks as compared with the other Bush Elephants, but how far this 
may have been due to the process of selecting largest tusks and elimi- 
nating these animals in ivory hunting is not clear. "In the days of 
van Riebeck (1653) elephant were plentiful as far south as the Cape 
Peninsula," but by the beginning of the next century seem to have 
become rare, for according to Theal the last one shot in this region 
was killed "just beyond Cape Flats in 1702; the expedition of Cap- 
tain Hop, in 1761, found plenty just north of the Oliphant River 
in what is now the district of Clanwilliam, while in the eastern half 
of the Colony, elephant hunting was regularly pursued till about 
1830. ... In Natal a few survived till 1860; in the north the 
hunters of the early part of the century made large bags near 
Kuruman ; Harris in 1836 shot chiefly in Magaliesberg of the western 
Transvaal; Gordon Gumming in 1846 in Sechele's country in 
northern Bechuanaland, and Livingstone and Baldwin, in 1849 and 
1858, found elephants innumerable on the Botletli River and near 
Lake Ngami, and finally Selous' hunting ground in the seventies 
and early eighties was in what is now Matabeleland and Mashona- 
land." Elephants were formerly so plentiful in the southeastern part 
of the Cape that an important ivory market was established in 1824 
at Fort Willshire. After 1860, however, the herds in the Knysna 
Forest and the Addo Bush were placed under government protection. 
The last elephant in Zululand was said to have been a solitary bull, 


which was killed in February, 1916, and its skeleton is now mounted 
in the Natal Museum. In Matabeleland, elephants existed in large 
numbers in 1872 and had been little hunted, but in the few succeeding 
years Lo Bengula's hunters in addition to Europeans swarmed into 
the region and in three years took out an estimated 100,000 pounds 
of ivory. Even then, tusks over 70 pounds in weight were rare, and 
the average was 40 to 50 pounds, rather small as compared with 
those farther north. By 1902 elephants had disappeared from the 
Transvaal, but in late years a few have come back into Kruger Park 
from adjacent areas of Portuguese East Africa. Shortridge sums 
up the present situation in the Cape Province and adjacent terri- 
tory: "Scattered and comparatively small herds of elephant still 
wander in Ngamiland, Southern Rhodesia, Portuguese East Africa. 
... In the Cape Province, the remnant of a herd is preserved in 
the Addo Bush. There may still be half a dozen or so in the Knysna 
Forest." In South-West Africa, there may be from 600 to 1,000 head 
in the Kaokoveld, but larger estimates are probably unwarranted. 
The other region where they occur is in the Caprivi, where two fair- 
sized herds are said to survive, one near the Kwando River, the 
other between Popa and Kagera. There are still elephants in south- 
western Angola and especially along the Kwando in the southeastern 
part. An estimate of the elephant population of any district is not 
easy to make, however, since on account of their wandering habits 
the same animals may appear within a short time at points far apart. 

In South Africa, aside from the restricted herds of Caprivi and the 
Kaokoveld, there exist four other herds: (1) that in the Knysna 
Forest, said to number, in 1935 about a dozen animals, which are 
under Government protection; (2) the Addo Bush herd, near Port 
Elizabeth, numbering, in 1933, about 16; (3) the Kruger Park herd, 
which seems most favorably situated and is believed to receive 
occasional increments from animals seeking this sanctuary from 
adjacent Portuguese territory; and (4) a small number that occa- 
sionally appear in straggling parties from across the Limpopo in 
times of drought and enter the northern Transvaal. Concerning 
the Addo Bush herd, in the early part of 1920 its numbers were 126, 
more than could well be maintained there, and so by Government 
order 110 were killed, and the remnant was confined to a more limited 
space, which apparently the animals more or less recognize and keep 
within its limits. A boring to supply them with water has been made 
to help in keeping them within these bounds, but the difficulty of 
restricting their wanderings is not easily overcome. In Kruger 
National Park there are said to be (1933) approximately 150-200 
elephants, in five separate groups. They tend to spread out from 
their fastnesses among the reed beds of the Letaba River (1934) . 

Apart from its great interest, from both esthetic and zoological 


points of view, the African Elephant has for centuries supplied a 
large part of the world's demand for ivory; it is a source of meat 
for many native tribes; and in recent years it has again been the 
subject of attempts at domestication, in this instance in the Belgian 
Congo, where imported mahouts from India as well as tame Indian 
Elephants have succeeded in rendering the Forest Elephant more or 
less tractable. Their timidity, however, often impairs their useful- 
ness, while the large amounts of food they require add to the diffi- 
culty of an economical value. According to Lavauden (1933, pp. 
21-22), in 1921, ivory to the amount of 800 tons of elephants' tusks 
was sent to the world's markets; in 1925 this had fallen to 500 tons, 
but the average weight of the tusks had considerably decreased as 
well. Elephants, on the other hand, often do much damage to the 
crops of the agricultural natives, wrecking their fields, granaries, and 
even houses at times. This damage, although "it is very doubtful if 
it would amount to 1 percent of the entire crop," is nevertheless at 
times a considerable loss, and of late years measures have been taken 
in countries under British rule to cope with this, by appointing an 
official to undertake elephant control through killing a certain num- 
ber in areas where they are reported to be doing such damage. In 
his book Elephant, David E. Blunt (1933), who had charge of this 
work in East Africa, reports that elephants seem very quickly to 
learn the bounds of regions to which they must be confined, and 
after a few of a marauding herd are shot the trouble to plantations 
is stopped for at least the time being. Thus while it is possible by 
this means to reduce greatly the elephant damage in agricultural 
areas near large forests or other country inhabited by herds of these 
animals, it is likely that with increase of settlement this protection 
of crops will become less needed, and the animals will gradually give 
way. Nevertheless there will undoubtedly be plenty of elephants in 
some sections of Africa for many years to come, in spite of hunting. 
Moreover, these will prove an asset on account of the returns from 
purchases of big-game licenses and additional fees for each elephant. 
In East Africa animals with tusks under fifty pounds in weight (the 
two together) may not legally be killed under penalty and confis- 
cation of the ivory. This limit, according to Brocklehurst (1933), 
has been lately reduced in Abyssinia from 30 to 20 pounds so that 
females now are killed. 

It appears from statistics that Uganda is likely to be one of the 
regions where elephants will long hold out and may be an asset in 
the way mentioned. In 1929, the Game Department reported a kill 
of 1,439 elephants, of which 1,135 were accounted for by the Govern- 
ment control operations. In 1931 the Game staff killed 1,211; in 
1933, the number was 1,380, and yet "with the exception of the Toro 
district, the southern portion of West Nile, and possibly the 
Mubende district, there is no reason to believe that elephant num- 


bers are other than steadily increasing in all parts of the protec- 
torate in which this great beast occurs." In 1934, it is said that 
2,716 elephants were killed in Tanganyika Territory. Taylor, in 
East Africa, July 9, 1936, believes that the method of control adopted 
is "the most humane method possible of enabling men and elephant 
to live in peace and concord in one territory." With the stopping 
of such methods of slaughter as once were practiced by natives in 
encircling elephant herds by grass fires and using pitfalls, and with 
the reduction of poaching for ivory, and the licensing of hunters, no 
doubt the hazards for the species are sufficiently lessened to counter- 
balance the large numbers just noted that are killed in control 
measures. The ivory from such elephants as are killed in this way 
is Government property and a source of revenue. Ivory is also a 
regular product of the Belgian Congo, where many animals must 
annually be killed, although at the present time this requires special 

For the future, the opinion of those conversant with the situation 
seems to be that in South Africa the relatively small areas of national 
reserves may continue to hold elephant herds indefinitely, but the 
size of the herds must be regulated by the area of the reserve and 
its suitability to their needs. With reduction to small numbers there 
is always a danger of an unlooked-for change which may be un- 
favorable. In South-West Africa, the numbers yet remaining are 
under government protection, so far as it may be enforceable, but 
the elephants here doubtless owe their continuation quite as much 
to the inaccessibility of their habitat. In the less settled parts of 
East Africa, they will continue in numbers and with the present 
efficient supervision of the game departments should prove on the 
whole a decided asset and attraction, notwithstanding a certain 
amount of local damage to plantations. In Uganda, where the herds 
are still abundant, there is evidence of slight increase in numbers in 
some districts, while in the great papyrus swamps of "the Sudd" of 
the upper Nile, they are present in great numbers and are likely to 
find this a safe retreat. Airplane photographs taken by the late Mar- 
tin Johnson in this region show some astonishingly large herds. The 
game warden of Uganda in his report for 1925 believes that with 
the spread of settlement and development elephants will have to be 
killed out or "expelled" from certain areas, but that, since extermi- 
nation is impossible and impracticable, good sanctuaries are neces- 
sary, which shall protect the main breeding areas of the herds. 
Elephants quickly learn to recognize the areas in which they are 
free from molestation, so that this trait will help to keep them within 
such bounds. A proper sanctuary, however, must include sheltered 
valleys with abundance of food and sufficient water, else at periods 
of drought the animals will move off in search of better localities. 

G. M. A. 


Order PERISSODACTYLA: Odd-toed Ungulates 

Family EQUIDAE: Horses, Zebras, and Asses 

Some conservative zoologists recognize but one genus in this 
family, while granting subgeneric status to the Horses (Equus), 
Zebras (Dolichohippus, Hippotigris) , and Asses (Asinus) . Others 
raise these subgenera to generic rank, and Shortridge (1934, vol. 1, 
p. 397) proposes an additional genus (Quagga) for the Quagga and 
Burchell's Zebra. Dr. Allen maintains a conservative viewpoint and 
employs the generic name Equus for all the Zebras (including the 
Quagga), while I prefer to keep both the African and the Asiatic 
Asses in a separate genus, Asinus. The single surviving species of 
Wild Horse (Equus przewalskii) is now confined to Mongolia. The 
nine forms of Zebras (two extinct) occupy eastern and southern 
Africa. One extinct and two living forms of African Wild Asses 
(Asinus atlanticus and A. africanus subspp.), with ranges in the 
northern and eastern portions of that continent, are herein recog- 
nized; also six forms of Asiatic Wild Asses (Asinus hemionus) , 
ranging from Mongolia and Tibet to Syria. The generally pre- 
carious status of the family is indicated by the fact that all but 
one of the Asiatic forms and all but six of the African forms are 
treated in the following pages. 

Przewalski's Horse; Mongolian Wild Horse; Mongolian Tarpan 


Equus Przewalskii Poliakov, Izviestiia Imper. Russk. Geogr. Obshchestvo, 
vol. 17, p. 1, 1881. (The type specimen was obtained by a "hunting- 
expedition sent by M. Tihonof from the post Zaisan to the sand deserts 
of Central Asia" (Poliakof, 1881, p. 19). Type locality restricted by 
Harper (1940, p. 195) to the oasis of Gashim, eastern Dzungaria (approxi- 
mately lat. 44 30' N., long. 90 E.).) 

SYNONYM: Equus hagenbecki Matschie (1903). 

Fios.: Poliakov, op. cit., pi. 1; Przewalski, 1883, pi. facing p. 40; Lydekker, 
1901, p. 284, fig. 65; Salensky, 1902, pi. 1, pp. 12, 16, 17, figs. 2-4; Proc. 
Zool. Soc. London 1902, vol. 1, pi. 13; Matschie, 1903, p. 582, fig.; 
Ridgeway, 1905, p. 27, fig. 18, p. 29, fig. 19; Wrangel, 1908, vol. 1, p. 3, 
fig. 1; Lydekker, 1916, vol. 5, p. 8, fig. 4; Peake, 1933, pi. 31, fig. a; 
Pocock, 1937, p. 715, fig.; Reed and Lucas, 1937, p. 129, fig. 44; Schmidt, 
1938, pi. 10. 

A very special interest attaches to this animal, as the only truly 
wild horse surviving in the world today. There is a remarkable 
dearth of first-hand information concerning it, especially during 
the past quarter of a century or so. Only one of the numerous 
scientific expeditions to Central Asia during recent years seems to 
have come into contact with it. It is somewhat doubtful if the 
alleged Mongolian Tarpans now exhibited in American zoos are 
purebred animals. 


Przewalski's Horse is distinguished from other horses by its erect 
mane and lack of a forelock. The following description is derived 
from Salensky (1902, pp. 7-18), who had more than a dozen speci- 
mens at his disposal, rather than from Poliakof (1881), who had 
only one. 

This species is of the size of a small ordinary horse; grown male 
with a height at the rump of 1,240 mm.; head relatively larger than 
that of the Wild Ass ; average length of ears, 140 mm. ; mane erect, 
highest (160-200 mm.) in the middle of its length; tail long, reach- 
ing in some individuals nearly to the hoofs, and provided on the 
dorsal side toward the base with short hairs, elsewhere with long 
hairs; hoofs rounder than those of the Kiang and the Kulan; 
"chestnuts" on all four limbs. 

Winter pelage lighter than that of summer; yellowish on the back, 
becoming lighter on the sides and almost white on the under parts. 
Summer pelage much shorter than that of winter, smooth, not wavy; 
back and sides light reddish brown, gradually changing to yellowish 
white on the belly; head colored like the back, but white on the 
muzzle about the nostrils and on the lips; ears light brown basally, 
darker at the tips; inner surface of ears white. Pronounced tufts of 
hair on sides of head in winter, and along entire lower part of head 
in summer. Mane dark brown, with shorter tufts of light gray hairs 
on each side; a median dorsal stripe of reddish brown, about 5 mm. 
wide, and distinctly visible only in summer pelage, extending along 
the entire back and on to the tail; a brown or black shoulder stripe, 
more noticeable in summer than in winter ; lower part of limbs more 
or less black (occasionally gray in younger animals) ; a black ring, 
up to 80 mm. wide, bordering the hoofs; inner side of legs gray, 
generally with distinct bars, up to the knees. 

The principal range seems to have been on both sides of the Altai 
Mountains in western Mongolia and in Dzungaria. But Prejevalsky 
(1876, vol. 2, p. 170) also reported Wild Horses much farther south, 
in western Koko Nor and in southeastern Chinese Turkestan : "Wild 
horses, called by the Mongols dzerlik-adu, are rare in Western 
Tsaidam, but more numerous near Lob-nor. They are generally in 
large herds, very shy, and when frightened continue their flight for 
days, not returning to the same place for a year or two. Their colour 
is uniformly bay, with black tails and long manes hanging down to 
the ground. [This last expression is, of course, wholly erroneous as 
applied to the manes.] They are never hunted, owing to the diffi- 
culties of the chase." This report, apparently based upon native 
information, does not seem to have been substantiated by later 
records, and is open to question. In this connection, however, it may 
be recalled that Sven Hedin has remarked (1903, vol. 1, p. 357) on 


the increasing scarcity of even the Wild Camel east of Lob-nor, 
owing perhaps to the increasing desiccation of the region. 

The animal must have been somewhat rare in the Altai region 
even before the advent of Europeans. Atkinson (1858), who made 
extensive explorations in that region about the middle of the past 
century, and comments frequently on the other large animals, does 
not refer to any personal encounter with the Wild Horse. However, 
in a later work (I860, p. 325), he describes the Kirghiz method of 
hunting "wild horses, which at this season [May] are found in 
great herds near the foot of the mountains" beyond the Hi River, 
apparently toward Issyk Kul. But his description of these horses 
as "varying in colour from black, bay, grey and white" creates 
considerable doubt as to whether they were truly wild or merely 
feral. Possibly the herds were composed of a mixture of both kinds 
of animals. Atkinson gives the Kirghiz name for the wild horse 
as "muss." 

Brehm (1876, p. 339) received a report of a second kind of Wild 
Horse (besides the Kulan), called "Surtake," which was said to 
occur about 250 versts southeast of the boundary post of Zaisan, in 
the Kanabo area. It was described as light yellow, with many light 
spots and a shorter tail than the Kulan's. 

Younghusband, referring to the region about the southern base of 
the eastern Altai, at about long. 96-100E., says (1888, p. 495) : 
"We . . . saw here . . . wild horses too the Equus Prejevalskii 
roaming about these great open plains." 

Ten years after Przewalski's discovery, the brothers Grum-Grshi- 
mailo took some specimens in 1889 at the oasis of Gashun, northeast 
of Guchen in eastern Dzungaria (Wrangel, 1908, vol. 1, p. 2) . 

The following report of Grum-Grshimailo (in Morgan, 1891, pp. 
217-218) probably refers chiefly to the Gashun area: "Springs 
enable the numerous animals inhabiting Dzungaria to exist; of 
these the most interesting is Prejevalsky's horse. . . . Prejevalsky 
himself, though he crossed the desert of Dzungaria in three several 
directions, never came across any of these wild horses, and if he 
wrote otherwise he was mistaking kulans he had seen in the distance 
for wild horses, a mistake the most experienced hunters are liable 
to make, for at that distance it is almost impossible to distinguish 
between them. . . . We were the first Europeans who, for twenty 
days, made a study of these interesting animals, adding the skins 
of three handsome stallions and one mare to our collection." 

Salensky (1902, pp. 2-3) records specimens from the following 
localities, chiefly in or near the Dzungarian Gobi: Gashun; the 
Kobdo region; behind the Baitik-Bogdo (Charamelechetai) ; between 
Nursu and Simigendse; Ebi Spring, near the Kobdo-Barkul route; 
Guchen Lake; and the River Bulunga. He gives the range (p. 63) as 


extending north to the Urungu River and Kobdo ; east to longitude 
90-91 E.; south to latitude 46 N.; and west to longitude 84 E. 
[=86?]. (This range is too restricted on the east and south.) 

In 1899 three newborn foals were captured and in the following 
year were brought to the estate of Herr Falz-Fein in Ascania Nova, 
southern Russia (Salensky, 1902, p. 20) . 

In 1901 Carl Hagenbeck sent a large expedition to Dzungaria for 
Wild Horses. His animals were caught in three different districts 
lying south of the Mongolian city of Kobdo. In the west the area 
consists of a wide plain, bordered on the east by the Altai Mountains. 
It is bordered on the north by the Kui-Kuius River, and on the 
south by the Urungu River, both of which rise in the Altai and 
discharge into the Tusgul [Ulungur?] Lake. This lake forms the 
western boundary of the plain. The second area is a plain which 
lies about 322 km. south of Kobdo and is enclosed by the Altai 
Mountains. The third group comes from the vicinity of Zagan Nor 
[apparently near long. 95 E.]. Foals from the three groups differ 
in color characters, though quite alike in general appearance 
(Wrangel, 1908, vol. 1, pp. 2-3) . 

The foals are dropped between the end of April and May 20. Their 
capture takes place as follows. Hundred of Mongols lie in ambush 
behind hills. As soon as they see a considerable number of mares 
and foals together, they rush upon them with loud cries. Since the 
foals can not keep up with the fleeing mares, the Mongols soon catch 
them with nooses on long poles. They are then conducted to near-by 
corrals, where Mongolian mares are ready to take over the duties 
of foster-mothers. Of the animals thus captured by the Hagen- 
beck expedition, 28 arrived in Hamburg in 1901 (Wrangel, 1908, 
vol. 1, p. 4) . 

"There is no doubt that the wild-horse . . . also inhabits the 
northern portions of that region [ Dzungaria]. We were never lucky 
enough to see any, but the natives, both Kalmuk and Kazak, all 
told the same tale, often volunteering the information that, in 
addition to the kulon, there were wild-horses. . . . They said, the 
meat was not so good [as the kulon 's]. They told us that there 
were large herds of them in the vicinity of Lake Ulungur, and east- 
wards along the southern foot of the Altai; also north of that range." 
(J. H. Miller, in Carruthers, 1913, p. 608.) 

From a point on the north side of the Altai, about 100 miles west 
of Ikhe Bogdo, R. C. Andrews reports (1926, p. 322) : "The wild 
. . . horses were two hundred miles to the southwest, they [some 
Chinese caravan men] said, just above the border of Chinese 

Morden writes (1927, p. 286) concerning a place in eastern Dzun- 
garia, northeast of Kucheng: "Around the spring, which our men 



we heard that the wild horses of 
. . were sometimes seen by cara- 

said was called Kainar Bulak . . 
western Mongolia and Dzungaria 
vans approaching the place." 

The range of the species was extended eastward by Lattimorc 
(1929, p. 228). In a journey of 1926, he mentions passing the "Yeh- 
ma Ching" or Wild Horse Well, which is situated in the Khara Gobi 

FIG. 33. Mongolian Wild Horse (Equus przewalskii) . After photograph 

in Brehm. 

west of Edsin Gol, at about latitude 42 N., longitude 98 E. "They 
say that on this fringe of the Khara Gobi there are wild horses 
(equ,us prjevalskii] and wild asses." This recalls Ridgeway's state- 
ment (1905, p. 28) : "Mr. Hagenbeck informs me that wild horses 
of another variety are said to exist 600 miles south of Kobdo. 
that is, somewhere in the great Gobi desert." 

Teichman also passed by the Wild Horse Well, in 1935, and makes 
the following remarks (1937, pp. 74-75) : "The Hardt-Citroen expe- 
dition followed from Suchou to Mingshui a camel trail which took 
them through this range. They found a region of rich pastures, 
abundant water, the haunt of ... the wild horse and wild ass. 
. . . Wild horses and wild camels are said to exist in this neighbour- 


hood. We saw no direct evidence of either. The wild ass is common 
and is often, with characteristic Chinese lack of accuracy, referred 
to as Yeh Ma ('wild horse') , which may explain the name of Yeh-ma 

Reymond, the zoologist of the expedition mentioned by Teichman, 
records (1932, pp. 807-809) a Wild Horse seen in May, 1931, and a 
carcass recently devoured by Wolves near the northern border of In- 
ner Mongolia at longitude 105 30' E., latitude "40" [ =42] N. The 
skull of the latter was identified by Professor E. Bourdelle of the 
Paris Museum. Reymond also heard that a favorite haunt of Wild 
Horses was the plateau of Pei Chan, occupying the extreme western 
triangle of Inner Mongolia. Other members of the Haardt-Audouin- 
Dubreuil Expedition (as it is here called) saw in 1931 two solitary 
animals in this general region: one in June, 20 km. west of Hou 
Hung Chuan (long. 96 E., lat. "40" [ = 42] N.), and one in 
December near Hsin Hsin Chia on the Kansu-Sinkiang frontier. 
The first of Reymond 's records is by far the easternmost one to date. 

"Przewalski's wild horse is found in small herds in Chinese Tur- 
kestan (Sinkiang) and Western Mongolia. It does not appear to be 
at all numerous, and should be protected if possible, if only because 
it is the sole surviving true wild horse in the world to-day. It is too 
small to be of any economic value, the so-called Mongol pony used 
by the Mongols and other Central Asian people easily supplanting 
it. The latter is probably a cross between Przewalski's horse and 
various domestic breeds, and is sufficiently hardy to live in the great 
wastes of Central Asia in a feral state." (Sowerby, 1937, p. 250.) 

Antonius writes (1938, pp. 558-559) : 

The statements of the brothers Grum-Grshimajlo and the expeditions of 
Falz-Fein and Hagenbeck for obtaining living specimens make it possible 
to give the geographic distribution about 1900. There was only one district on 
the Northern ranges of the Ektag Altai: the neighbourhood of the Zedsig- 
Nor, called also Zagan-Nor, and three or four in the deserts on the southern 
ranges: the steppes on the upper Urungu, the Ebi-mountain, the Gashium- 
desert. If my Russian information is correct and I have, alas! no doubt 
that it is, the Przevalski horse has been extirpated since the great war and 
the Russian revolution, the old fork-muskets of the Mongolian hunters 
having been replaced by modern fire-arms of great power. Therefore it is 
probable that the descendants of the Falz-Fein and Hagenbeck-imports living 
in Ascania Nova, Woburn Abbey, and in a few Zoos in Europe, America, and 
Australia, are the last survivors of the Przevalski-horse, and of the true wild 
horse in general. 

There would appear to be considerable likelihood that Przewalski's 
Horse, if not exterminated outright, has proceeded far along the road 
to extinction through dilution with the ponies of the Mongols. In 
Salensky (1902, p. 21) we find a report of domestic mares mating 
with wild stallions. In remarking on the variations in color exhibited 


by Przewalski's Horse in different parts of its range, Lydekker says 
(1912, p. 89) : 'These differences suggest that there has been some 
admixture with domesticated breeds." Who can even say that the 
type specimen was purebred? 

The chances of mixture with domestic stock are suggested by 
Carruther's report (1913, pp. 532-533) of "immense droves of horses 
running half wild over the prairies" in the vicinity of Barkul, 
southern Dzungaria. "We . . . believe that the real 'wild animals' of 
the Barkul basin signify the great herds of unridable horses which 
roam untamed over the steppes. These form an Imperial Stud, and 
are said to number fifteen thousand, the pick of which are trans- 
ported yearly to Pekin." 

Domestication. On this subject Peake (1933, pp. 99-100) says: 

There can be no doubt that the horse was first tamed in the grasslands 
of Central Asia, for it is only there that the wild horse, known as Przewal- 
sky's horse, is to be found to-day. The first mention of the horse that has 
been met with is in a document from Babylon, dating from before 2000 B. C., 
in which it is called the ass from the East. This indicates the direction from 
which it came, but it does not seem to have been introduced into Mesopotamia 
before the arrival of the Kassite conquerors about 1746 B. C. We have, 
however, some reasons for believing that it had been tamed at an earlier 
date. Into the north of Mesopotamia there had arrived some centuries earlier 
a people known as the Kharians, some of whom were later called the Mitanni. 
These, we know, were great horsemen. They occupied the country around 
Haran, which lies between the Tigris and the Euphrates just below the points 
at which they emerge from the mountains, and they seem to have arrived in 
that district from the North-east, probably from the Persian plateau, whence 
later the Kassites descended upon Mesopotamia. The horse was well known 
also to the Hittites, the capital of whose kingdom lay in the centre of Asia 
Minor. These people are believed to have arrived there about 1900 B. C. 
from the North-west. All this evidence tends to show that the horse was 
used as a means of transport both in Persia and upon the Russian steppe 
well before 2000 B. C. It seems likely that it was first tamed in that part 
of the world, or still farther east in Mongolia, as early as 3000 B.C., if not 
before that date. 

Lamm Wild Horse 

EQUUS sp. 

Surprising news of a generally overlooked and probably extinct 
Wild Horse in the Kolyma Basin of northeastern Siberia is fur- 
nished by Pfizenmayer (1939, pp. 112-113). While excavating the 
frozen carcass of a Mammoth on the Beresovka River in 1901, he 
questioned two Lamut visitors as to 

what sort of wild animals they found in their distant hunting-ground on the 
Omolon. To our great astonishment Taitshin mentioned the wild horse. As 
zoologists thought wild horses existed only on the steppes of central Asia, we 
received his statement very doubtfully, though Amuksan confirmed it by a 
quite professional imitation of horses neighing. The reliability of the natives 


is such that we did not imagine they were spinning a yarn when they told us 
about the wild horse in the tundras bordering the forests of this vast area. 
They described in detail its size equal to that of a Yakut horse its long 
whitish -grey hair, and its flesh, which was very fat and pleasant to taste. 
If the description were really that of wild [= feral] horses, it was a puzzle 
how and when their tame ancestors could have reached this quite uninhabited 
Arctic region. And if they actually existed in the district between the two 
largest tributaries of the Kolyma the Omolon and the Anjui which had 
never yet been explored by any scientist, it was a very interesting matter 
which scientists would find it well worth while to investigate. 

Pfizenmayer's assumption that these horses were descended from 
tame ancestors is by no means necessarily correct. In this connection 
it is of interest to recall Hay's opinion (1913, p. 9) that in the Yukon 
Basin and adjacent parts of Alaska horses "became extinct about 
the middle of the glacial epoch." 

Pfizenmayer writes further (pp. 176-177) : 

The prehistoric wild horse to which is probably related the animal that 
Przevalski, the Russian explorer of Asia, discovered in 1870 and called a 
wild horse has left remains everywhere in central and northern Siberia. 
There is hardly one place on the banks of rivers and lakes in the district 
of Yakutsk in which prehistoric remains of animals have been found that 
has not yielded skeletal fragments of the prehistoric wild horse. 

In the landslide on the Beresovka we found, among the debris between 
the larch trunks lying around in confusion and the masses of fallen earth, the 
perfectly preserved upper skull of a prehistoric horse, to which fragments of 
muscular fibre still adhered. . . . 

An exiled student told me, in Verkhoyansk on my way back from Kolymsk, 
that an ivory hunter had found the carcass of a horse four years before, 
sticking half out of the frozen earth in a fissure in the bank of a lake in the 
tundra, in the northern part of the district. According to the description 
by the man, who puzzled over the find unusual there the parts of the 
body sticking out of the ground showed a covering of very long greyish-white 
hair. Certainly the Yakut horse, a vigorous breed of pony, with probably a 
strain of the wild horse, also has long hair to protect it from cold. Since, 
however, there were no Yakut settlements for hundreds of miles round the 
site of the find, we may conclude that the body was that of a prehistoric 
wild horse. But it was naturally neither investigated nor salvaged. 

Determination of the relationship between this prehistoric horse 
and the Recent Lamut Wild Horse must await the acquisition of 
suitable museum material. 

In commenting upon some earlier publication of Pfizenmayer's 
findings, Antonius writes (1938, p. 559) : "One might suppose that 
these white horses are descendants of any semiferal Jakute-breed 
the Jakutes being the most northern horse-breeders, but ... it, 
could be possible that these wild horses of the Lamutes are the last 
survivors of a northern branch of the Caballus-Group, and there 
are some indications for a formerly much greater distribution includ- 
ing not only Eastern Siberia, but also Alaska. Since the excavation 
of the Beresovka-Mammoth there are no records of the Lamute 


The plausibility of a white horse on the Siberian tundra is en- 
hanced by Janikowski's account of present-day domesticated descen- 
dants of the Wild Horse of Poland. He refers (1942, p. 682, figs. 4-5) 
to two survivors "which had the remarkable and unique property of 
turning white in winter .... Every winter they changed the 
mouse-grey summer coat . . . into a snow-white coat, only the face, 
fetlocks, mane and tail retaining the dark colour." 

European Wild Horses 

EQUUS spp. 

The taxonomic and nomenclatural status of European Wild Horses, 
especially during the more recent historical times, becomes extraor- 
dinarily complicated owing, on the one hand, to the lack of ade- 
quate material and authoritative data and, on the other hand, to 
the probability of interbreeding with domestic types. The technical 
nomenclature is too involved to be discussed in detail in this brief 
account. Opinions differ as to whether some of the described forms 
were truly wild or were mixed with the blood of domestic horses. 
Only purebred wild animals come properly within the scope of the 
present report. 

Remains of Pleistocene or older horses have been recorded in 
various localities from India and Turkestan to Spain, France, and 
England ; some of these were doubtless ancestral to the present-day 

In classical times Strabo reported Wild Horses in Spain. In the 
Middle Ages there are records of Wild Horses in Germany, Poland, 
Lithuania, and Russia; but there is some question as whether all 
of these records refer to purebred wild animals. 

In 1768 S. G. Gmelin collected four Tarpans in the Government 
of Voronesh, Russia. Pallas (1811) reported Wild Horses as inhab- 
iting the steppe country from the Dnieper to the Altai and beyond 
into Central Asia, but as partly mixed with feral animals. Hamilton 
Smith (1845-1846, pp. 160-166, pi. 3) received information from 
Cossacks and others early in that century concerning truly wild 
animals in Russian Turkestan and Mongolia. 

Antonius (1912, p. 513) mentions three animals captured alive in 
Russia as late as the period 1853-66 ; he considers these the last Wild 
Horses taken in Europe. However, Lydekker (1912, p. 81) suspects 
that even Gmelin's specimens were hybrid Tarpans, and it is all 
the more to be doubted that the animals of 1853-66 were purebred. 
Antonius (1912, p. 516) has given the name of Equus gmelini to the 
three last-mentioned animals, at the same time stating that Gmelin's 
specimens were probably though not certainly identical with them. 
If the specimens on which the name Equus gmelini was based were 


not purebred, the name can hardly be applied to their truly wild 
forebears in Russia, which have been extinct for probably more than 
a century. In the present rather chaotic state of the nomenclature, 
I feel unable to fix upon any one of the numerous names proposed as 
applying strictly and validly to the form represented by the last 
truly wild and purebred horses of Europe. 

A few quotations from the literature will indicate some of the 
varying opinions on a complex subject. The later accounts can 
hardly refer to purebred wild animals. Poliakof (1881, p. 20) says: 

The information regarding the tarpan collected by Rytchkof, Gmelin, 
Georgi, and Pallas is of so contradictory and confusing a nature that many 
zoologists have decided that the so-called wild horses, or "tarpans," were 
not, strictly speaking, wild, but tamed horses which had resumed their 
wild state on recovering their liberty .... Pallas . . . assumed the feral horses 
. . . roaming over the steppes of the Yaik [Ural] and the Don as well as on 
that of Baraba to have originated from domesticated horses owned by Kirghiz, 
Kalmuks, or other wandering tribes, and to have become wild. . . . Un- 
fortunately we have no reliable information on this legendary tarpan since 
the end of the last century, not a single traveller either in Siberia or Russia 
having communicated any information concerning it during the present 

"The nearest approach to truly wild horses existing at present are 
the so-called Tarpans, which occur in the steppe-country north of 
the Sea of Azoff, between the river Dnieper and the Caspian. They 
are described as being of small size, dun color, with short mane and 
rounded, obtuse nose. There is no evidence to prove whether they 
are really wild ... or feral." (Flower, 1892, p. 83.) 

Calinescu (1931, p. 82) reports Equus caballus gmelini as sur- 
viving in Moldavia, Rumania, up to 1716. 

Vetulani (1933, pp. 281-282) gives the following account for 
Poland. Hacquet (1794) describes wild horses kept in a zoological 
garden near Samosch. They increased to such an extent that some 
were shot and others were sent to Lemberg for use in combats with 
carnivores. In Kajetan Kozmian's reminiscences of the years 1780- 
1815 (published in 1858), we read likewise of wild horses in a 
zoological garden near Zamosc or Samosch. They were allowed to 
become extinct, apparently because in winter it was necessary to 
provide barns and hay for them. These two references concern the 
last wild Forest Tarpans of Poland ("E. c. gmelini ssp. silvatica") . 
From Brincken (1874) we learn that this stock was derived from 
the last wild horses in the Forest of Bialowies, and that finally they 
were captured in the zoological garden near Samosch and divided 
among the peasants. This represented the last stage in the domesti- 
cation of the European Wild Tarpan. We still find in this vicinity 
representatives of the Forest Tarpan type in an especially pure and 


typical form. It is proposed to introduce and preserve some of them 
in the Polish National Park at Bialowies. 

"According to Vetulani, the enigmatic wild white horse described 
by Herodotus as grazing in the northern marshy land may well have 
been the Polish wild pony grazing in the Polesie bogs situated close 
to the Bielowieza Forest" ( Janikowski, 1942, p. 682) . 

According to Niezabitowski (1934, p. 196), E. gmelini Antonius 
lived formerly in the steppe region of eastern Poland, while E. 
gmelini silvaticus Vetulani inhabited the Bialowies Forest up to 
the middle of the eighteenth century. 

Heptner reports (1934, pp. 431-433) that the last example of 
Tarpan was seen in 1914-18. It lived at that time on an estate in 
Dubrowka, Mirgorod district, Government of Poltava, and was 
very old. It had been purchased as a young animal from German 
colonists, who shortly before had destroyed a small herd of wild 
horses. Hitherto the last Tarpans in South Russia were supposed to 
have died out in the 1870's. They survived longest on the steppes 
of the Government of Cherson. 

From the foregoing it may be gathered that it is virtually impos- 
sible to state even approximately when the last truly wild repre- 
sentatives of the genus Equus perished in the various European 
countries. Even the names that should be applied to them are far 
from settled. The type of Equus caballus caballus Linnaeus is the 
Scandinavian domesticated horse of the time of Linnaeus obviously 
at least subspecifically distinct from the Russian Wild Horse. 
Certainly all Wild Horses are now extinct, with the exception of 
Przewalski's Horse of Central Asia. 

For a fuller account of the Wild Tarpan and its relations, Lydekker 
(1912, pp. 71-116) may be consulted. For a discussion of some of the 
nomenclatural problems involved, see Harper (1940, pp. 195-197). 

Antonius (1938, pp. 557-558) gives the following illuminating 
account of the caballus-group of horses : 

In times not long before the beginning of historical days there were true 
wild horses of the Caballus type spread over the whole Eurasiatic continent 
from the Atlantic to the Pacific, and from the shores of Northern Siberia 
to the Indian Ocean. Only two, or at the utmost three, of the many local 
and geological races have survived until our days. The first of these was a 
mouse-dun horse, which Albertus Magnus, the great interpreter of Aristotle, 
means when he calls the colour of the wild horse "cinereus," i. e., ash-coloured. 
There can be little doubt that these ash-dun or mouse-dun wild horses were 
often intermingled with escaped domestic horses of the feral breeds, thence 
spread over Europe. But there are some indications by which in many cases 
their true wild nature may be ascertained. The one is the high value of 
these horses for princely gifts, the other the short upright mane, and the 
third the uniformly ash colour, so often recorded. If the first Duke of 
Prussia, Albert von Hohenzollern, sent wild horses as highly esteemed gifts 
to the mightiest sovereign of his days, the Emperor, and also to the Arch- 


duke Ferdinand and others, there can be no doubt that these horses were a 
truly royal game like the Urus and Bison and not of a little valued feral 
breed. And if the mouse-colour and the short mane are recorded for some 
of the last survivors in Poland and Southern Russia, it must be almost 
certain that there was at least a strong strain of true wild blood in these 
horses. S. G. Gmelin, one of the many German explorers of Russia in the 
days of the great Catherine, hunted these "Tarpans" in 1763 in the surround- 
ings of Bobrowsk, Woronesh. After him the author named these horses 
scientifically "Equus gmelini" but perhaps there is an earlier name: Equus 
silvestris v. Brincken, dedicated to the mouse-dun wild horses of Poland, 
surviving in the forest of Bialowieza until the middle of the eighteenth 
century, and in another game park until 1812. Although protected very 
strictly against poaching and illegal hunting, the wild horse in Prussia vanished 
in the second half of the sixteenth century. In the well-known forest of 
Bialowieza, Poland, the "Tarpanis" were hunted as royal game in 1409, 
when King Wladislaw Jagiello arranged a great chase in honour of his cousin, 
Witold of Lithuania. In the immense forests they survived until the eighteenth 
century, when they were extirpated before the time of the famous hunting 
of the Saxon Kings. Their last refuge in the Poland of to-day was accord- 
ing to Vetulani the great game park of the Count Zamoyski, situated at 
Zwierzyniec, near Bilgoraj. Here they were strictly protected, until in a 
severe winter between 1810 and 1820, probably from 1812 to 1813, the feeding 
was impossible. The last survivors were captured and given to the peasants 
of the surrounding country. According to these facts there are in no other 
district of Poland more typical "Tarpans" among the little horses of the 
peasants than in the surroundings of Bilgoraj [c/. Janikowski, 1942]. Ve- 
tulani has proved these Polish wild horses as a more or less degenerated 
branch according to their being adapted to the unsuitable wood life of 
the Eastern or Russian Tarpan. 

The latter vanished from the fertile country of Woronesh before 1800, but 
survived on the steppes of Tauria and Cherson until the middle of the nine- 
teenth century. The last herds were certainly more or less intermingled with 
feral horses, but the short mane being recorded even for the last example, 
demonstrated the predominance of true wild blood. F. von Falz-Fein, the 
well-known founder and owner of the matchless Ascania Nova Zoo, has 
told the life-history of that last wild horse of Europe, an one-eyed old mare, 
lingering for years around the feral horses of a certain Durilin, covered by 
a domestic hehorse, captured, escaped with its filly, and some years later 
hunted and killed on the ice by the peasants of Agaiman. 

There is only one drawing from a living example hitherto known: in the 
description of the travels of Gmelin, edited by Pallas after the tragic death 
of his comrade. This picture, drawn by Borisow from a one year old mare, 
was later on copied by Schreber in his "Naturgeschichte der Saugetiere." 

It must be recorded that experiments for the rebreeding of the mouse- 
dun Tarpan were started both in Germany and in Poland. In the Schorfheide 
near Berlin and in the Munich Zoo the Germans try the rebreeding by 
crossing the true yellow-dun Mongolian wild horse with mouse-dun mares 
of various domestic breeds, while in Bialowieza the Poles settled upon some 
most typical descendants of the last Bilgoraj wild horses, selected out of a 
great number of peasant-horses in that district, without any interbreeding 
of strange blood. The question is, which of the two trials will have the 
better results. 

The home of the mouse-dun Tarpan extended eastward over the river 
Don and probably to the right bank of the Wolga. It is possible in earlier 


days that these horses were also spread over the Caspian steppes, but as it is 
difficult to distinguish the different records about other Equidae, e. g., the 
Kulan, it is impossble to confirm that opinion. 

The hillier steppe-country between the Wolga and the Ural-Mountains, 
in the days of Pallas already crossed by a line of Kossak-posts, were roamed by 
another wild horse. Pallas gives in his great travel-work, the well-drawn 
portrait of a young filly, captured in the surroundings of Tozk then a little 
Kossak post. That picture resembles in a high degree the Przevalski-fillies, 
imported by Hagenbeck in 1899 and 1900. Together with the statements of 
Pallas about the colour (Isabella to light bay), the "suberect" mane, the 
tail, etc., there can be no doubt that these horses were almost as pure-bred 
wild horses of the yellow-dun Przevalski-type as ever roamed the Dzungarian 
Gobi. In the time of Pallas the wild horses were spread in scattered troops, 
more or less intermingled with escaped domestic horses, over the steppes of 
Western Siberia. Georgi, one of Pallas's fellow-workers, reports that they 
were extirpated by a desolating horse-sickness in 1785 which destroyed also 
the herds of the Kirghises and Kossaks, causing the death of about 85,000 
horses. In 1876 the species was rediscovered by the great Russian explorer 
Przevalski in the Dzungarian steppes south of Kobdo and named after him 
by Poljakoff "Equus przevalskii." 



Equus quagga Gmelin, Linnaeus' Syst. Nat., eel. 13, vol. 1, pt. 1, p. 213, 1788. 
(South Africa.) 

FIGS.: G. Edwards, Gleanings of Natural History, p. 29, pi. 223 (col.), 1758; 
Ridgeway, Proc. Zool. Soc. London 1909, pp. 563-586, text-figs. 157-180, 
reproductions of early figures and photographs of preserved specimens. 

The vernacular name, Quagga, of this handsome zebralike species 
is said to be derived from the Hottentot khoua khoua, in imitation 
of its barklike cry. The Boers, however, often applied the same name 
to the BurchelFs type of zebra, and it is sometimes loosely used for 
that animal by writers. 

Harris, whose folio Portraits of the Game and Wild Animals of 
Southern Africa, 1840, provides some first-hand information on this 
species, wrote that it stands 4.5 feet at the withers and has a total 
length of 8.5 feet, but Cuvier (quoted by W. L. Sclater, 1900, vol. 1, 
p. 295) gives the height at the shoulder as slightly less, about 4 feet 
1 inch. The general ground color of head, neck, and body was dark 
rufous brown or bay, becoming gradually more fulvous and fading 
off to white behind and beneath. The midline of the back was 
marked by a broad dark stripe. Against the background of bay, the 
forehead was marked with longitudinal stripes and the cheeks with 
narrow transverse stripes of buff, "forming linear triangular figures 
between the eyes and the mouth." Muzzle black; neck and anterior 
half of the body banded and brindled with creamy brown, broader 
and more regular on the neck (extending across the short erect 
mane), but becoming finally lost in spots and blotches on the rear 



half of the trunk. The legs, tail, and under surfaces were white, 
with sometimes a short midventral dark line, and usually a black 
spot behind the fetlocks. Individuals appear to have varied con- 
siderably in the width and extent of the paler stripes and in the 
amount of whitish on the rump, tail, and belly. These differences 
have been made the basis of several subspecific names, but it is 
now agreed that they are best considered as only variations of a single 
species. Although Pocock earlier believed that the Quagga was 
merely a southernmost form of the BurchelPs Zebra, and that certain 

FIG. 34. Quagga (Equus qwagga). After Standard Natural History. 

individuals nearly bridged the gap between the extremes, it has since 
been shown that the two are doubtless separate species, and that the 
Quagga, in addition to the well-marked color characters, was further 
distinguished by cranial differences as well. For according to 
Schwarz (1912b) the skull is the smallest of the three South African 
species of zebras, and is characterized by its relatively wide zygo- 
mata, narrower bony eye ring, broader forehead, greater separation 
of the temporal ridges, the presence of a small suborbital pit, and 
by having the posterior border of the nasals heart-shaped. While 
such characters may be subject to individual variation, they may for 
the present be regarded as valid. 

The Quagga seems first to have been brought to the notice of natu- 
ralists by George Edwards, who in his Gleanings (1758) published 


a colored plate of a female, which he supposed to be the female of the 
Mountain Zebra. Buffon in 1782 (Hist. Nat., Suppl, vol. 6, p. 85) 
was the first to give an authentic account of the animal on the basis 
of notes supplied him by Allamand, from Colonel Gordon, a South 
African resident. Subsequently Edwards' description became the 
basis of Gmelin's name Equus quagga. So far as the records show, 
the Quagga always had a somewhat restricted range and was con- 
fined chiefly to the southeastern corner of the Cape region, from 
Algoa Bay westward at least to Prince Albert (where Barrow reports 
it in 1801), and Swellendam, some 100 miles east of Cape Town, 
where Sparrman first saw it, northward to the Orange Free State and 
the Vaal River, and coastwise to the Kei River. There is practically 
no record of travel in South Africa between the days of Kolben in 
1705 and the visit of the Swedish naturalist Sparrman in 1775. The 
latter, however, found the Quagga as near the coast of Algoa Bay as 
Uitenhage. He secured a foal which is still preserved in the Riksmu- 
seum at Stockholm. Quaggas were apparently still plentiful in the 
first quarter of the last century, especially in the districts of Aberdeen 
(Lichtenstein, in 1804) and in Fraserburg and Hanover where Bur- 
chell in 1812 found them abundant, in troops of 30 to 50 on the plains. 
These made an impressive sight traveling in single file as was their 
curious habit, or when startled, wheeling in unison like a squad of 
cavalry. It was said that they frequently associated on the plains 
with the White-tailed Gnu or with ostriches; whereas the Burchell's 
Zebras preferred the companionship of the Brindled Gnu. Where, 
as in the Orange Free State, the range of the Quagga met or over- 
lapped that of Burchell's Zebra, it is said the two did not mingle. 
The Boer farmers evidently took heavy toll of them in these years, 
greatly reducing their numbers, and using the meat to feed their 
workers although themselves preferring more tasty kinds of game. 
The result of this constant persecution was that by the late 50's the 
Quagga was practically extinct south of the Orange River. According 
to Bryden (1889) the last known instance was of two shot in 1858 
near the Tygerberg, a solitary mountain rising abruptly from the 
plains near Aberdeen. His informant, the successful hunter, recalled 
the affair well. Farther north, however, in the Orange Free State, 
Quaggas were still numerous. About 1865, the Boers of this state 
began the exploitation of the large game of the region for their hides. 
With characteristic industry and deadly skill they gathered and 
shipped to the coast hides by the wagonload, among which those of 
Quagga and Burchell's Zebra were especially in demand. They also 
made use of Quagga hides for grain sacks, and Bryden (1889) men- 
tions seeing old Quagga-skin sacks still in use at the time of his visit. 
The exact date of the final extermination of the Quagga is unknown, 
but it is generally believed that the species continued well into the 


'seventies in the Orange Free State, probably, according to W. L. 
Sclater (1900) , "till 1878 at least," but he adds, "it is difficult to 
obtain any accurate information on the subject, as in so many cases 
this and BurchelPs Zebra are confused together, especially as they 
were both known under the name of quagga." 

In the earlier days of the last century and even shortly before, 
Quaggas were occasionally tamed and also exported alive to the 
zoological gardens of Europe. In disposition it was said to be much 
more tractable than the BurchelPs Zebra, in captivity quickly be- 
coming docile and tamable. On various occasions they were broken 
to harness, and Sir William Jardine even mentions that a Mr. Sheriff 
Parkins early in the nineteenth century drove a pair in London, 
and was often seen in Hyde Park riding in a phaeton after them. 
Probably one of the first Quaggas to reach Europe alive was the one 
belonging to His Royal Highness the Prince of Wales, from which 
in 1751 George Edwards made his colored drawing. The specimen 
now in the Paris Museum was brought to the menagerie of the King, 
at Versailles, in 1793. Others were later imported by animal dealers 
such as Frank at Amsterdam. Of the various specimens extant in the 
museums of Europe, the larger part were brought in alive and 
received by the museums after having died in captivity. Thus the 
locality of capture is in most of these cases unrecorded. In 1858, Sir 
George Grey presented to the Zoological Society of London a male 
Quagga which died six years later, in 1864. "It is the mounted skin, 
skull and skeleton of this male which is now in the British Museum" 
(Ridgeway, 1909). Previously in 1851 the Society had purchased 
a female Quagga which survived in Regent's Park, until 1872, ap- 
parently nearly the last living example of the species of which any 
positive record exists. Further, this was the only living Quagga ever 
to be photographed, and the picture has been reproduced by Ly- 
dekker in his Guide to the Specimens of the Horse Family and by 
Ridgeway in his paper of 1909. The skin was not in condition to be 
preserved; but it is said that the skeleton was saved and mounted, 
although at the present time it has been lost sight of and is evidently 
not the one now in the British Museum. Finally, the last known 
living specimen seems to have been one that died in the Berlin 
Zoological Garden in 1875. The skin is mounted in the Zoological 
Museum in that city, and the skeleton is also preserved there. 

Combining the lists of Ridgeway (1909) and Hilzheimer (1912), 
the known specimens of the Quagga in the museums of the Old World 
are the following (arranged alphabetically by location) : 

1. Amsterdam Museum. Mounted specimen, and separate skull. Figured 
by Lydekker (Proc. Zool. Soc. London, 1904, vol. 1, p. 430, text-fig. 86) and 
by Ridgeway (1909, p. 579, text-fig. 170). 

2. Basle Natural History Museum. A mounted female from Silo (Shiloh), 



Cape Colony, presented in 1864. Figured by Ridgeway (1909, p. 565, text- 
fig. 157). 

3. Berlin, Zoological Museum. Mounted female, that died in the Berlin 
Zoological Gardens in 1875; also its skeleton and two other skulls. Figured by 
Ridgeway (1909, p. 578, text-fig. 168). 

4. Cape Town Museum, South Africa. Mounted foal, from Beaufort West, 
about 1860. Figured by Ridgeway (1909, 580, text-fig. 171). 

5. Darmstadt Museum. Mounted specimen. 

6. Edinburgh, Royal Scottish Museum. A mounted specimen, purchased by 
the University of Edinburgh during the year ending June 1818. Figured 
by Ridgeway (1909, p. 575, text-fig. 165). 

7. Elgin Museum, Scotland. A mounted head and neck, from King Wil- 
liam's Town, 1861. Figured by Ridgeway (1909, p. 581, text-fig. 172). 

8. Frankfurt a.M., Senckenberg Museum. A well-mounted skin and its 
skull, received in 1831 by exchange with the Leiden Museum. 

9. Leiden, Dutch State Museum of Natural History. Mounted male and 
its skeleton, shot near Steenbergen, June 15, 1827. Figured by Ridgeway 
(1909, p. 577, text-fig. 166). 

10. London, British Museum. Ridgeway (1909, p. 574) has cleared away 
the confusion regarding the number and origin of the specimens in this 
Museum. Apparently the only one is a male skin, mounted, and the skeleton 
of the same animal, which had been presented to the Zoological Society of 
London by Sir George Grey in 1858 and lived in the Society's Gardens until 
its death in 1864. The female Quagga which lived in the Gardens from 1851 
to 1872, was photographed in life, but its skin, upon its death, was in too poor a 
state to be preserved. Its skeleton, however, was mounted but cannot now 
be traced. The male specimen as mounted is figured by Ridgeway (1909, 
p. 573, text-fig. 163) and the living female is figured by Ridgeway (1909, p. 575, 
text-fig. 164) from York's photograph of the animal. 

11. Mainz Museum, Germany. According to Hilzheimer (1912) there were 
four mounted Quaggas in this collection, but Schwarz (1912) who also examined 
them, asserts that one of the four is a Burchell's Zebra. 

12. Munich Natural History Museum. A mounted specimen purchased in 
1835, and a separate skull that may or may not belong to the same individual. 
It was this specimen that was the original of the figure by Wagner in "Schreber's 
Saugthiere, Supplement." Figured by Ridgeway (1909, p. 579, text-fig. 169). 

13. Paris Museum of Natural History. A mounted specimen. According to 
the communication of Dr. E. L. Trouessart, it was received living after the 
institution of the Museum's menagerie from the old menagerie of the King, 
at Versailles in 1793. No more precise locality is given for it than "Cape of 
Good Hope." Figured by Ridgeway (1909, p. 577, text-fig. 167). 

14. Stockholm, Riksmuseum. The mounted specimen here is a full-grown 
fetus, brought back by the Swedish traveller Sparrman in 1775. It is therefore 
the oldest extant specimen and appears to have the pale stripes clearer and 
extending farther back than usual. While the exact locality is unrecorded, 
Sparrman mentions that he first saw Quaggas at Swellendam. Figured by 
Ridgeway (1909, pp. 570, 571, text-figs. 160, 161), both from a photograph and 
from a recent painting. 

15. Stuttgart Museum, Germany. According to Hilzheimer (1912) this 
museum contains a skull and footbones of the Quagga. He further mentions 
that in the Stuttgart Altertums Collection is a miniature model of a Quagga, 
of which he gives a figure, and suggests that it was probably prepared from 
the two animals which Frederick I had in his menagerie in 1812-16. It may 
therefore have a certain authenticity. 


16. Tring Museum, England. A mounted specimen, interesting for the 
distinctness of the posterior stripes. It was described and figured by P. L. 
Sclater (Proc. Zool. Soc. London, 1901, vol. 1, p. 166) who believed that it 
was the same as the animal formerly living in the London Zoological Gardens 
from 1851 to 1872. That this, however, is not the case was made clear by 
Ridge way, who publishes a letter from E. Gerrard (who sold the specimen 
to Lord Rothschild) stating that he had purchased the animal as an old 
mounted specimen from a Mr. Frank of Amsterdam and had remounted it 
before selling it to the Tring Museum. Figured by Ridgeway (1909, p. 569, 
text-fig. 159). 

17. Turin, Zoological Museum. A mounted female, and its skull. The 
specimen was purchased in 1827 from the English dealers, Leadbeater father 
and son. It was made the type of Equus trouessarti, figured and described 
by Camerano (1908, pi.). 

18. Vienna Museum. A mounted female, procured by Ecklon, in 1836. 
The specimen was described by Lorenz (Proc. Zool. Soc. London, 1902, vol. 1, 
p. 32) and figured from a photograph which is again reproduced by Ridgeway 
(1909, p. 568, text-fig. 158). 

19. Wiesbaden Museum, Germany. A mounted specimen, male, which was 
bought in 1865 from Frank, the Amsterdam dealer. It has no more definite 
locality than "South Africa." Figured by Ridgeway (1909, p. 572, text-fig. 162). 

From this enumeration it appears that there are in the museums of 
Europe 17 mounted skins (one a fetus) , a mounted head, 3 skeletons, 
and 7 skulls ; while elsewhere the only known specimen is a mounted 
skin of a foal, in the South African Museum. 

G. M. A. 

Burchell's Zebra or Bontequagga 


Asinus burchellii Gray, Zool. Journ., vol. 1, p. 247, 1824. (Little Klibbolikhoni 

Fontein, Bechuanaland, South Africa.) 
FIGS.: Gray, op. tit., pi. 9, figs. 1, 2; Lyon, M. W., Jr., Proc. U. S. Nat. Mus., 

vol. 32, pis. 1-3, 1907 (photographs of mounted specimen) ; Cabrera, Jour. 

Mamm., vol. 17, p. 97, figs. 1-5 (pattern diagrams), 1936; Pocock, Proc. 

Zool. Soc. London, vol. 1, p. 485, fig. 48, 1909 (photo.). 

Although, on account of its somewhat variable pattern, the Bur- 
chell's Zebra has been divided into many nominal races, only four of 
these are regarded as valid by Cabrera (1936) in his recent review. 
Of these four, the typical burchellii is now extinct, and there are few 
specimens preserved. 

About the size of a small horse, with erect mane and tufted tail, 
the color pattern consists of alternating dark-brown or black stripes 
and whitish stripes. Of the former, there are about ten on the neck, 
a vertical shoulder stripe, four body stripes, then on the flanks three 
or four that turn back dorsally, somewhat paralleling the median 
black stripe. The characteristic feature of typical Burchell's Zebra 
is that the lower haunches and both fore and hind legs lack the small 



transverse stripes that are increasingly developed in the more 
northern races, and the tail is white. 

The former range of this race was rather limited and covered what 
is now the Orange Free State and southern Bechuanaland, in South 
Africa, but apparently did not extend to the south of the Orange 
River. Over the plains of this region it once abounded "in countless 
thousands," but with the coming of white hunters, followed by 
settlers in the Orange River colony, it had already become rare by 
the middle of last century. Many were exported to Europe for 

FIG. 35. Burchell's Zebra (Equus burchellii burchellii). After Brehm. 

zoological gardens, and it is in part from these that have come the 
few specimens still preserved in museums. There is a specimen in the 
British Museum, one in the Tring Museum, and a third in the 
Bristol Museum, in England, and there is a mounted one in the U. S. 
National Museum, and one in the Paris Museum, with a few others in 
other museums, as Berlin, Leiden, and South Africa. The last 
living specimen, so far as known, was one kept in the London 
Zoological Gardens, where it was received apparently in 1909, after 
evidently having been in captivity for a period. 

From Benguela west to Southern Rhodesia and Zululand, this race 
is replaced by the race antiquorum, with more cross-striping on the 
upper parts of the limbs. At the present time this animal still exists 
in some numbers in the west of South-West Africa. From the Lim- 
popo River northeastward to the Loangwa and Rovuma Rivers is 


found the race selousii, distinguished by "having the limbs striped to 
the coronet, and the body with numerous narrow stripes and few and 
faint shadow stripes." Still farther northeastward, is the race bohmi, 
inhabiting the plains country of eastern Africa north to Lake Rudolf, 
in which the shadow stripes (between the clear black stripes) are 
absent and the haunch stripes broad and black, and the limbs cross- 
striped nearly to the hoof. This race is still common. 

The chief enemy of the zebras, apart from man, is the Lion, which 
seems specially fond of zebra meat and finds it easily obtainable. 
The zebras go in herds which may at times be of large size. Fre- 
quently Gnus, of one species or another, associate with these herds, 
as if for companionship. Zebras have at times been tamed and 
trained to harness but are of rather uncertain disposition and no 
great use of them in this way has been made. Their meat, though 
relished by the natives, is not popular among white hunters, accord- 
ing to Selous. The hide is often used as leather. In regions where 
agriculture is practiced, Zebras often become a nuisance to the 
ranchers by stampeding and breaking through barbed-wire fencing. 
They are said to have a good deal of curiosity, and especially where 
mules or donkeys are in camp, will often approach closely to survey 
them. According to Major Flower, they often in captivity live over 
12 years, and have been known to reach 28 or 29 years in zoological 
gardens. The name "quagga" is said to be derived from the noise 
they make, a sort of honking bark. They are much dependent on 
water, and drink at least once a day, often at night, but approach 
the waterhole with caution, for fear of lurking lions. 

Although, with the exception of the typical burchellii, none of the 
races is in present danger of extinction, their numbers will undoubt- 
edly diminish except in areas where large extents of grasslands 
as in East Africa afford them range. 

G. M. A. 

Mountain Zebra. Wildepaard (Boer). Dauw (Hottentot) 


Equus zebra Linnaeus, Systema Naturae, ed. 10, vol. 1, p. 74, 1758. (Probably 
the Drakensberg and other mountain ranges, Cape of Good Hope.) 

Hartmann's Mountain Zebra 


Equus hartmannae Matschie, Sitzber. Ges. Naturf. Freunde Berlin, p. 174, 
1898. (Between Hoanib and Unilab Rivers, South-West Africa.) 

FIGS.: Of typical form: Lydekker, 1912, pi. 20, fig. 1; J. E. Gray, Knowsley 
Menagerie, pi. 56, 1850; Pycraft, 1936, p. 850, fig. Of E. z. hartmannae: 
Haagner, 1920, fig. 66; Zukowsky, 1924, fig. 6; Maydon, 1932, pi. 125; 
Shortridge, 1934, vol. 1, pi. opp. p. 389; Pocock, 1937, p. 717, fig. 


Since there is still some doubt (Shortridge, 1934, vol. 1, p. 389) 
whether the Mountain Zebra of the dry South-West African uplands 
is recognizably distinct from the typical E. zebra, the two may be 
treated together, pending further study of specimens. Also some 
authors, among them Captain Shortridge (op. cit.), prefer to regard 
this as a genus distinct from Equus or from the South African 
Quagga, calling it Hippotigris. Since in skeletal and tooth characters 
it is very little different from the horses, a conservative course is to 
regard the latter as a subgenus of Equus. 

The Mountain Zebra was the first of the zebras known to 
Europeans and is the smallest of the three species, standing about 
12 hands high (48 inches) at the shoulder. Length of head and body 
7 feet 4 inches; tail, with terminal hairs, 23 inches (W. L. Sclater, 
1900). Sclater gives the following description: 

Body, head, and limbs closely covered with black or almost black stripes, 
broader than their white interspaces; on the face the dark markings below 
the eyes become reddish passing into large nostril patches of the same color, 
but the muzzle itself is black; . . . ears long and rather narrow, posteriorly 
the basal two-thirds striped, the terminal third black, the extreme point 
white; . . . longitudinal dorsal stripe only noticeable over the haunches, 
transverse stripes of the barrel extending back over the haunches to the 
base of the tail forming here the so-called gridiron pattern; no shadow 
stripes; hairs along the back to the shoulders reversed; belly white, except 
for a longitudinal dark band running along its anterior portion which is 
never reached by the transverse body stripes; limbs transversely marked 
down to the hoofs, . . . the pasterns being quite black; . . . hoof rather 
narrow, compact and solid; tail reaching the hocks with a median black line 
and traces of transverse bars at the base; the distal quarter with a tuft of 
long black hairs. 

Hartmann's Zebra is believed to differ from the typical race in 
its larger size and more widely spaced stripes, so that the pale stripes 
are equal to or even slightly wider than the black ones. The legs 
"are almost evenly banded black and buff the black not predomi- 
nating as in zebra." However, this pattern varies individually and as 
yet it is uncertain whether the characters claimed are relatively 

As its name implies, this zebra was an upland species, living in 
the mountains, "from Great Namaqualand (and possibly Damara- 
land), through the various ranges of Cape Colony to the Great 
Drakensberg chain, and thence to the end of that range. ... At 
the present day," wrote Bryden (1899, p. 94) , "it is only to be found 
in small troops here and there in Cape Colony. It is very doubtful 
whether any now remain in Great Namaqualand, where, sixty years 
ago, Sir James Alexander found them in considerable numbers. It is 
probable that the Hottentots . . . , who are excellent shots . . . , 
have destroyed the last remnants ... in ... Great Namaqualand. 



In Cape Colony, where these zebras are, as far as possible, preserved, 
small troops are to be found in the mountains of the Sneeuwberg, 
Witteberg, Tandtjesberg, Zwartberg, the Winterhoek, and one or two 
other ranges. A few still linger along the Drakensberg. . . . Near 
Cradock, . . . only a few years since, a troop of twenty was seen." 
Under date of January 23, 1935, Herbert Lang, the well-known 
explorer and authority on large game mammals of South Africa, 

FIG. 36. Mountain Zebra (Equus zebra zebra). After photographs in 
Brehm and Newnes. 

wrote that "there must be still about a hundred Mountain Zebras 
in various places in spite of the reports to the contrary. In 1926 
I traveled through all these regions to observe these zebras in their 
haunts. One must have seen these herds of Mountain Zebras on 
their actual trails to be enthusiastic about them and their protec- 
tion. There can hardly be anything more fascinating." By 1937, 
according to editorial notes in the Journal of the Society for the 
Preservation of the Fauna of the Empire, a census of these zebras 
showed the following 45: in the Oudtshoorn Area, near George, 20 
on the farm of Peter. Heyns ; in the Cradock Area, 10 on the farm of 
Paul Michau, 8 on the farm of Lombard, 7 on that of Osborn. The 


proportion of sexes among these animals was believed to be very 
uneven with "a great shortage of mares." This number is still less 
than a few years previously, for E. L. Gill, writing in December 1932, 
said that at Cradock there were "two or three small herds which seem 
likely to die out. The Oudtshoorn herd on the farm Mount Hope, 
has been carefully preserved by the owners, the brothers Heyns, and 
is still flourishing. It numbers somewhere about 70 animals and con- 
stitutes the chief hope for the survival of the species." 

Efforts have been made at various times in recent years to induce 
the Government to purchase a portion of the Mount Hope Farm as a 
permanent Mountain Zebra reserve. In 1937, the Government at 
last voted to ask the Parliament of the Union of South Africa for 
7,600 for the establishment of such a reserve and hopes are high that 
it may actually be created, before it is too late. Notwithstanding 
that the species is protected at all times by the South African 
Government, the actual enforcement of this protection has in the 
past been difficult. It is one of the species listed for complete pro- 
tection by the London Convention of 1933. 

While "the advance of civilization" is blamed for the reduction 
in numbers of this zebra, no doubt much blame must also be laid 
upon the native and white poachers with modern rifles, as well as 
to other methods of extermination. Bryden (1899) wrote that occa- 
sionally the weather is so "severe among the Cape mountains that 
even the tough zebra succumbs" and that in "the old days in Cape 
Colony, the Boers were in the habit of hunting these animals for the 
sake of their hides and of capturing the young alive for the purpose 
of being broken to harness." For in the last century "a fashion for 
using Mountain Zebras in harness seems ... to have sprung up in 
the Mauritius, and ... a good many of these animals were exported 
from the Cape to meet the requirements of the French colonists. 
... A premium of 20 was at the same time offered for the young 
of these animals delivered in Cape Town." "The Boers, to save 
themselves the trouble of shooting, occasionally succeeded in driving 
a number of these animals over the edge of a precipice, thus securing 
the skins at their leisure" (Bryden, 1899) . 

Although Bryden (1899) feared that within the "next fifty years 
this zebra will have joined the ranks of extinct creatures," there 
seems still some hope of preserving a remnant, owing chiefly to the 
interest of those farm owners on whose lands the survivors still hold 
out, and an awakened enthusiasm on the part of the Government to 
do what it can. 

While the future of the Mountain Zebra in the Cape Province is 
none too rosy, it still occurs in the form hartmannae in small numbers 
among the mountain ranges of the western and northwestern parts 
of South-West Africa and northward across the Cunene into south- 


western Angola at least as far as Elephant Bay, 100 miles north of 
Mossamedes, where, however, it is not found more than 30 miles 
inland. Shortridge (1934, vol. 1, pp. 390-396) has gathered together 
the available information concerning it in this region. The eastern 
limits are found in the Kaokoveld about a hundred miles from the 
west coast. Here it is sometimes found in association with the Bonte- 
quagga (E. burchellii antiquorum) but is much fewer in numbers. It 
is partial to the crests of arid gorges, and its small cupped hoofs 
are adapted for rough country. Large numbers are said to be found 
in the Omaruru and the Maltahohe districts, but elsewhere they are 
less common. Steinhardt saw them digging for water in sandy river 
beds of this arid country, making pits half a meter deep. They may 
not drink regularly but sometimes keep away from water as long as 
three days. They are shy and suspicious and difficult to approach 
under usual conditions. The Cape Mountain Zebra is believed to be 
a slow breeder, with foal every second year or so. The period of 
gestation is said to be about twelve months. The height at the 
shoulder is said to be in hartmannae about 52 to 54.5 inches, hence 
somewhat taller than the typical race. 

G.M. A. 

Nubian Wild Ass. Nubischer Wildesel (Ger.) 


Asinus ajricanus Fitzinger, Naturgesch. Saugethiere, vol. 3, p. 667, 1857. 
(Lydekker (1916, vol. 5, p. 38) gives the type locality as "Nubia (accord- 
ing to Matschie, Erythraea) .") 

SYNONYM: ? 'Asinus asinus dianae Dollman (1935). 

FIGS.: Proc. Zool. Soc. London 1884, pi. 50, fig. 2, 1885; Lydekker, 1904, 
pi. 20, 1912, pi. 20, fig. 2, and 1916, vol. 5, p. 37, fig. 16; Antonius, 1929, 
p. 290, fig. 1; Zammarano, 1930, p. 87, fig. 

This subspecies is "by no means common" (Brocklehurst, 1931, 
p. 15). 

"General colour of upper-parts greyish-fawn, with the muzzle, a 
broad ring round each eye, . . . and the under-parts, white or 
whitish ; the legs being of the same pale hue, with some greyish on the 
front surface, and a few small dark spots on each side of the fetlocks. 
The mane ... is short, upright, and dark brown or blackish .... 
The narrow dorsal stripe ... is continued as a thin line well on to 
the tail"; the two branches of the shoulder stripe are about 5-6 
inches in length. "The long hairs of the terminal tail-tuft ... are 
mingled black and grey. The ears are about 10J in. in length, and 
are black at their tips .... On the inner side of the lower part of 
the fore-leg is a chestnut patch." Height of male at shoulder, 45^ to 
47i inches. (Lydekker, 1904, p. 594.) Baker (1867, p. 56) gives the 
height of a male from the Atbara River as 55-56 inches. 



A. a. dianae Dollman (1935), from south of Tokar near the 
Eritrean boundary, is so weakly differentiated from ajricanus as to 
seem scarcely worthy of recognition ; it may represent a slight inter- 
gradation toward somaliensis. 

"The Nubian wild ass ... inhabits , . . Sennar and Nubia, its 
range formerly extending as far as the fifth cataract of the Nile 

FIG. 37. Nubian Wild Ass (Asinus asinus ajricanus) 

.... Year by year the range of this race appears to become more 
restricted; and unless protective measures be taken, there is danger 
that it may be exterminated." (Lydekker, 1908, p. 66.) 

Heuglin (1861, p. 19) reports Wild Asses as occurring from Suakin 
to the Nile at Berber, in all northeastern Sennaar, and in the plains 
of the Barka River. He says he met with them commonly about 
the ruins of Wadi Safra, then on the Atbara, and along the route 
from Taka toward Suakin ; and during the rainy season they appear 
as far north as the Desert of Korosko. (Korosko is on the Egyptian 
part of the Nile, at about lat. 22 30' N.; but Flower remarks (1932, 
p. 432) : "There appear to be no certain records of genuine wild 
asses having occurred in Egypt during the nineteenth century.") 


Sir Samuel Baker (1867, pp. 55-56) writes concerning the Wild 
Ass along the Atbara River: "Those who have seen donkeys in 
their civilized state have no conception of the beauty of the wild 
and original animal. . . . The animal in its native desert is the per- 
fection of activity and courage; there is a high-bred tone in the 
deportment, a high-actioned step when it trots freely over the rocks 
and sand, with the speed of a horse when it gallops over the bound- 
less desert. No animal is more difficult of approach; and, although 
they are frequently captured by the Arabs, those taken are invari- 
ably the foals, which are ridden down by fast dromedaries, while 
the mothers escape." 

Matschie gives (1894, p. 73), as the northernmost locality, the 
Wadi el Homar, a little north of Berber. 

"Their flesh is eaten by the Arabs of the Soudan. They are 
ordinarily met with in twos and threes, or small herds." (Bryden, 
1899, p. 70.) 

"This animal is found at the foot of the Gebel Hennah, near 
Tokar. It is common in the Khor Sabbat parallel to the Khor 
Baraka. Captain O'Connor informs me that he has often seen them 
at the Khor Sabbat, on the plain of Tokar." (Anderson, in Ander- 
son and de Winton, 1902, p. 330.) 

"Neither the wild asses nor the zebras of Africa are pursued with 
much enthusiasm by sportsmen, and the first-named animals are 
so shy and wild that whilst it is very difficult to get within shot of 
them on foot, if they are hunted on horseback they are so fleet and 
enduring that they can only be overtaken with great difficulty even 
by a really fast horse. . . . Thus the wild ass is seldom shot, and 
is probably of less interest to the average sportsman than any other 
African game animal." (Selous, 1914, p. 36.) 

"The Wild Ass is found in the Sudan in the neighbourhood of the 
Atbara River in the provinces of Berber and Kassala; it is also 
found in the Red Sea Province south of Suakin. 

"They have been strictly protected for a number of years, and 
although by no means common there is not, at present, any danger 
of their being exterminated." (Brocklehurst, 1931, p. 15.) 

Some years before 1932 Wild Asses were fairly common in the 
region of the Baraka Wadi near the border of Eritrea (Maydon, 
1932, p. 203) . 

In all probability the very few Eritrean specimens now extant live 
in the region- of Upper Barca (De Beaux, 1935, p. 12) . 

Powell-Cotton (in Dollman, 1935, p. 134) writes concerning Wild 
Asses south of Tokar near the Eritrean boundary: "Beween 18 Feb- 
ruary and 2 March 1934 we saw the animals on four occasions, as 
follows, 1, 2, 2, 3, and secured the two specimens permitted us. 
. . . The Arabs . . . leave the Wild Ass unmolested as they do not 


eat its flesh, and they told us they were more numerous across the 
Eritrean boundary." 
Antonius (1938, p. 560) writes: 

[The Nubian Wild Ass] until recent times spread over the mountainous 
semi-deserts of Nubia and the Eastern Sudan from the Nile to the shores 
of the Red sea. The last specimens of Nubian origin known to European 
observers were two males shot by Sir Reginald Loder about 1925, near the 
Gebel Raboba, on the Erythrean frontier, and an old female, living in the 
Zoological Gardens at Rome for many years, the photograph of which is 
reproduced by Zammerano in his "Fauna e Caggia." To-day there is a living 
specimen in the New York Zoological Park, and also a stallion in the Zoo 
at Rome. The Nubian ass was domesticated by the ancient Egyptians at 
least in 3500 B. C., and being hunted by the King Rameses III. as is shown 
at the temple of Medinet Habu it was well known as a wild animal in 
Egypt until at least 1100 B. C. . : . 

The so-called "Nubian Asses" shown in many Zoos, are by no means pure 
bred descendants of the true wild stock; therefore it seems nearly certain 
that the Nubian ass is to-day as thoroughly extinct as his Atlantic cousin. 
[This remark on extinction is somewhat premature, in view of the statements 
just quoted from Brocklehurst, Maydon, De Beaux, and Powell-Cotton.] 

All forms of the Wild Ass in Africa enjoy complete protection 
under the London Convention of 1933. 

Domestication. According to Lydekker (1916, vol. 5, p. 37), the 
typical subspecies of the African Wild Ass (A. a. asinus) is repre- 
sented by the Domestic Ass of Asia. He also states (1912, p. 217) 
that the wild animal was first tamed in the Mediterranean countries. 
This ancestral wild animal was doubtless distinguishable from the 
present domesticated animal, but whether it was identical with any 
of the wild forms now recognized (africanus, atlanticus, or somali- 
ensis), it is impossible to say. Werth (1930, p. 351) suggests Abys- 
sinia as the place of first domestication. If, however, the wild ances- 
tor inhabited the eastern Mediterranean region, where no wild repre- 
sentative of Asinus asinus now exists, it may have differed from any 
of the currently recognized forms. In the account of Asinus hemionus 
hemippus (p. 368) I have suggested the possible occurrence of some 
wild form of Asinus asinus in the Palestine region within compara- 
tively recent times. 

While space does not suffice to discuss the subject of domestication 
at any length, the following quotation from Peake (1933, pp. 98-99) 
is apropos here: 

The ass was used at a very early date, both in Egypt and in Mesopotamia, 
but it is impossible to say at present to which region to ascribe the priority. 
Towards the close of the predynastic period in Egypt, just before 3400 B. C., 
the Libyan tribes, who dwelt in the desert to the west of the Nile Delta, 
possessed large herds of asses, and this indicates that this animal had been 
known to and possessed by them for a long time. At Ur, in Mesopotamia, 
asses were used by those kings and queens who were buried in the famous 
death-pits, accompanied by their slain retainers, and these must be relegated 


to quite as early a date, and are probably much earlier. All our evidence 
goes to show that asses had been tamed, and were used as beasts of burden, 
both in Egypt and Mesopotamia, between 4000 and 3000 B. C., and may have 
been domesticated at a considerably earlier date. 

On the island of Sokotra there are herds of wild or feral asses 
closely resembling A. a. africanus in color but of smaller size (stand- 
ing from 38 to 40 inches at the shoulder) . They are regarded as "the 
survivors of Nubian ancestors brought from the Red Sea coast by, 
probably, the ancient Egyptian incense collectors." Their introduc- 
tion is presumed to date back "for some thousands of years" per- 
haps a sufficient length of time for their insular habitat to have pro- 
duced degeneration in size. "As they are never shot at and rarely 
molested by the natives, they were by no means wild." (C/. H. 0. 
Forbes, 1903, pp. xxxviii, 6, 9-11, pi. 2.) 

[There has been much discussion as to the possible existence of 
Wild Asses in the Sahara. Tristram (1860, p. 318, as quoted by 
Blyth, 1862, pp. 363-364) "heard that wild Asses were to be occa- 
sionally found in the Soufa desert, on the route to Ghadames." He 
was shown one that "had been caught when very young, and was 
considered unusually tame for one of his species." It was "of a rich 
slatish ash-colour"; dorsal stripe and shoulder stripe present; nose 
and limbs white; mane and tail blackish. The adults were said to 
be "very difficult to entrap and impossible to train." 

On a journey through the Tuareg country in 1913-14 Geyr von 
Schweppenburg (1917, p. 298) learned that wild or feral asses were 
not uncommon there in previous years, but by that time had virtually 
disappeared, having been captured, shot, and fed to dogs. The local 
Mohammedans did not eat the flesh. He was inclined to consider 
that at least some of the asses were genuinely wild, and not merely 
feral. He also mentioned reports of Wild Asses in this region by 
previous travelers (Duveyrier, Bissuel, Benhazera, and others). 

According to Spatz (in Werth, 1930, p. 347) , these wild or feral 
asses constantly molested the domesticated animals and often led 
them astray. 

Antonius (1931, pp. 133-136, pi. 3, fig. 3) calls attention to some 
wild-living asses reported by Fraulein von Wagner-Jauregg in the 
Hoggar massif in the southern territories of Algeria, where the 
Tuaregs distinguish them by name from the Domestic Asses. A 
captured foal showed evidence of a strong wild-blooded component, 
even if it coud not be considered a pure-blooded wild ass. It is feared 
that these wild-living asses of the Hoggar will disappear if energetic 
protective measures are not adopted. 

On the other hand, Hilzheimer and Spatz (Zeitschr. fiir Sauge- 
tierk., vol. 17, p. 15, 1932) express the opinion that the animals of 
the Hoggar are merely Domestic Asses that have run wild. Selous 


(1914, p. 35) and Lavauden (1933, p. 20) state that Wild Asses do 
not pass west of the Nile. 
Antonius (1938, p. 560) says: 

It is very interesting that the only mention of a wild ass captured in the 
Western Sahara, given by Canon Tristram, agrees not with the appearance 
of the Atlantic, but with that of the Nubian, race. Although very exactly 
describing the coloration of his wild born ass, Tristram does not say any- 
thing about banded limbs. The wild asses once roaming in the neighbour- 
hood of the caravan route to Chadames [Ghadames] and Chat [Ghat] were 
therefore probably not of the strongly marked Atlanticus-type, but of the 
Nubian with unbanded limbs. This opinion is strengthened by the colours 
of the so-called "ahoulil" of the Tuareg wild or feral asses, lingering in the 
Ahaggar mountains in single pairs and small troops. A filly of these ahoulil, 
captured in 1927 in the heart of the Ahaggar mountains for the Schonbrunn 
Zoo, but dying before transported to Europe, was of exactly the same Nubian 
type. Probably there is in these ahoulil at least a strong strain of originally 
wild blood, more or less intermingled with the Atlanticus blood of escaped 
domestic Tuareg donkeys. 

Finally comes the extremely interesting information from Mai- 
brant (1936, p. 27) that in French Central Africa Wild Asses are 
restricted to the massif of Tibesti. Views differ as to their origin and 
systematic status, but Malbrant inclines toward the opinion that 
they are genuinely wild animals of the subspecies africanus. They 
exist in the region of Zouar and, farther north, in the Tarsoa, moun- 
tainous ridges situated north of Emi Koussi. Here they are not rare. 
The plateau of Daski and the region of Trotron (between Yebi and 
Zoumri) likewise shelter many. They live in bands of as many as 
30 or 40 individuals. The natives capture young ones in snares near 
the water-holes, train them, and use them as pack animals. 

A view differing from Malbrant's is held by Thesiger, who remarks 
(1939, p. 441) that in Tibesti "donkeys are extensively used, and 
many have run wild among the mountains probably for generations." 

The solution of the problem of the Tibesti Asses awaits the collec- 
tion of specimens.] 

Somali Wild Ass. Somali- Wildesel (Ger.) 


A[sinus] taeniopus var. Somaliensis Noack, Zool. Garten, vol. 25, no. 4, p. 101, 
1884. ("Somaliland"; type locality restricted by Lydekker (1916, vol. 5, 
p. 39) to "Berbera district of [British] Somaliland.") 

FIGS.: Nouv. Arch. Mus. Hist. Nat. Paris, vol. 5, Bull., pi. 5, 1869 (subsp.?) ; 
Proc. Zool. Soc. London 1884, pi. 50, fig. 1; Akeley, 1914, pp. 112, 115, 117, 
figs.; Zammarano, 1930, p. 88, fig.; Schmidt, 1938, pi. 11. 

In British Somaliland this Wild Ass "is strictly preserved, but . . . 
much reduced in number" (Antonius, 1938, p. 560) . 

It is more strongly built than A. a. africanus] general color a deli- 


cate reddish ash-gray ; snout gray ; behind it a broad, light gray band 
from the nose to beyond the corner of the mouth ; a light ring about 
the eye ; inner surface of ears ash-gray, with black border and tip ; 
outer surface of ears yellowish red ; mane light gray basally , fuscous 
above; shoulder stripe absent; a dark but not very pronounced 
median dorsal stripe extending from the lumbar region to the tail 
tuft; forelegs yellowish gray anteriorly, light gray posteriorly; dark 
bands about all the legs up to the level of the body, but only on the 
anterior side of the forelegs (Noack, 1884, pp. 101-102). Height at 
shoulder about 51 inches (Menges, 1887, p. 262). Pocock (1909, 
p. 528) remarks on a seasonal change of color, from clear gray in 
summer to sandy fawn in winter. 

Menges (1887, pp. 263-267) gives the following account: 

The range is apparently restricted to Somaliland and part of the 
Red Sea coastal plain south of Massaua, Eritrea; it probably in- 
cludes the Danakil region and extends south to the Webi Shebeli. 
A particular habitat is the coastal lowlands, where the animal is 
not exactly rare; another favored haunt is the barren Hekebo Pla- 
teau (2,000 feet high), southeast of Bulhar. It is commonly found in 
herds of 5 to 20 head, and is extremely shy and cautious. 

In general it leads a rather undisturbed existence, though occa- 
sionally falling victim to the Leopard or the Lion. Most of the 
Somalis do not touch the flesh, but one or two tribes pursue the 
animal to some extent. Only a few hides are brought from the 
interior to the coastal markets. On the other hand, in the coast 
districts of the Red Sea the Wild Ass is eagerly pursued, with the 
object of shipping captured animals to Arabia, where they are used 
for crossing with the Domestic Ass. One result of this crossing is 
the hardy and beautiful riding ass of Yemen. Among the Somalis 
themselves one finds many Domestic Asses bearing evidence of 
crossing with the wild animals. 

"In certain parts of Guban, notably in the sterile district lying 
near the coast, about twenty miles east of Berbera, the Wild Ass is 
not very uncommon. We met with it also in considerable numbers 
on the high plateau west of Laferug, and also saw some individuals 
south of the Golis Range .... The flesh of these animals is very 
good, almost the best we ate in Somaliland .... It does not seem 
to be a very plentiful species even in the country of its nativity, 
and I should judge it would not require much persecution to speedily 
extinguish the race." (Elliot, 1897, pp. 139-140.) 

"The Somali wild ass is fairly common. I first met with them 
about twenty miles to the south of Berbera, and they are also found 
on the plateau to the south of the Golis range. They do not live on 
the mountain ranges, but frequent the low stony hills in the desert. 


They go in small herds. The largest I saw consisted of five." (In- 
verarity, in Bryden, 1899, p. 71.) 

''They are common to the eastward of Berbera, behind Siyaro, in 
among the sand dunes and rocky hills, and also south of Bulhar in 
similar localities, especially around the Issitugan Valley. South of 
the Golis Range they inhabit the low stony hills around Halo, 
Haloka Yer, and near Segig; they are also found on Negegr Plateau." 
They are "usually seen in herds of four or five individuals and not 
uncommonly singly." (Drake-Brockman, 1910, p. 103.) 

Akeley (1914) gives an account of hunting Wild Asses about 30 
miles from Berbera. One reason for their scarcity is indicated in 
his statement (p. 117) that "one English 'sportsman' boasted of 
having killed twenty-eight." 

De Beaux (1928a, p. 6) records seven specimens from Italian 
Danakil. He also remarks (p. 13) that the present subspecies is sepa- 
rated from ajricanus by the Ethiopian plateau, which approaches the 
Red Sea at the Gulf of Zula [Annesley Bay]. 

Thesiger, who traversed Abyssinian and Italian Danakil in 1933, 
found Wild Asses quite common north and south of the lower 
Hawash (Neumann, 1935, p. 153) . 

Antonius (1938, p. 561) writes: 

Because many hides from Berbera, as well as from Danakil, although typical 
in all other points, show a more or less developed shoulder cross, there can 
be no doubt that neither the existence of it nor its absence is thoroughly 
typical [of somaliensis}. . . . 

It is, alas! to be feared that the Abyssinian war has its consequences for 
the African wild asses: warring soldiers, and especially askaris, are never 
the best protectors of vanishing game! Whether the "Asinus somaliensis" 
exists also in Southern Abyssinia or not is not positively known. A well- 
informed Austrian, who had been living in Abyssinia for many years, told me 
that he had seen wild asses in the Bale country on the upper Juba. I suggest 
for geographical reasons that the animals are not true wild asses, but either 
Zebras or domestic donkeys of a feral breed, similar to the beautiful asses 
of the Turkana people on the western shore of Lake Rudolf, at first seen by 
von Hoehnel and Count Teleky, and since recorded by modern visitors to that 

Lydekker (1916, vol. 5, p. 39) records a specimen from as far 
south as "Shebeli Valley, Somaliland." 

Atlas Wild Ass; Algerian Wild Ass 


Equus asinus atlanticus. P. Thomas, Mem. Soc. Ge"ol. France, ser. 3, vol. 3, 
no. 2, p. 45, 1884. (Recent Quaternary deposits, Oued Seguen, near 
Constantine, Algeria.) 

SYNONYM: Equus asinus atlanticus Werth (1930). 

FIGS.: Thomas, op. cit., pi. 2, figs. 7, 7a; Werth, 1930, p. 348, fig. 3; Jennison, 
1937, pi. facing p. 145. 


Apparently a Wild Ass inhabited Algeria up to at least about 
A. D. 300, but subsequently became extinct. It was probably distinct 
from any form now living. 

P. Thomas (1884, p. 45) based the name Equus asinus atlanticm 
upon a mandible with teeth, found in late Quaternary deposits near 
Constantine, Algeria. In these he found characters apparently inter- 
mediate between those of Pliocene Hipparion and those of the 
present-day Domestic Asses of Algeria. 

The name atlanticus may be applied at least provisionally to the 
Algerian Wild Ass of Roman times. Werth (1930, p. 350, map) 
indicates the presumable former distribution as including Morocco 
and Tunisia as well as Algeria. 

"In a Roman villa at Bona, in Algeria, was found a large and 
well-preserved picture, dating from about A. D. 300, of an African 
hunt. Its main effect is a representation of a drive of carnivora. . . . 
The use of the lasso is illustrated in the same picture, where a 
Numidian, riding bareback and stirrupless, is throwing one at a wild 
ass." (Jennison, 1937, pp. 145-146.) 

Antonius (1938, pp. 559-560) says: 

The true asses of African origin the wild stock from which our domestic 
donkey descends belong to the many mammalia which became totally extinct 
in our days. There were in Roman times at least three local races, one of 
which became extinct before it was ever seen by a modern zoologist. It was 
the "Asinus atlanticus Thomas," well known from the rock picture of Enfouss, 
Algeria, published erroneously as "Quagga" by Frobenius. An excellent Roman 
mosaic at Hippo Regius, the modern Bone, also shows that donkey. It 
possessed a well-developed shoulder stripe, strongly marked limbs, and the 
ears perhaps a little shorter than its East African cousins. The geographic 
distribution of these Atlantic asses seems not to have exceeded the ranges 
of the Atlas mountains. The time of their extinction is unknown. 

Mongolian Wild Ass; Chigetai; Dziggetai; Kulan ; Kulon 


Equus hemionus Pallas, Nov. Comm. Acad. Sci. Imper. Petropolitanae, vol. 19, 
p. 394, pi. 7, 1775. ("Ad Lacum Tarei Davuriae" = Tarei Nor, on the 
Siberian-Mongolian boundary, about lat. 50 N., long. 115 E.) 

SYNONYMS: Equus onager castaneus Lydekker (1904); Equus (Asinus) 
hemionus bedfordi Matschie (1911) ; Equus (Asinus) hemionus luteus 
Matschie (1911). (C/. Harper, 1940, pp. 197-198.) 

FIGS.: Pallas, op. cit., pi. 7, and 1781, pi. 1; Lydekker, 19046, pi. 27 (bedfordi) ; 
Lydekker, 1904c, pi. 18 (castaneus); Lydekker, 1912, pi. 15, fig. 2 
(castaneus); Lydekker, 1916, p. 13, fig. 6 (castaneus); Carruthers, 1913, 
pis. facing pp. 602, 606; R. C. Andrews, 1924, pp. 152-156, figs., and 1926, 
pi. facing p. 129. 

During recent years, in all its vast range, the Mongolian Wild 
Ass seems to have been reported as plentiful in only one region that 
about Orok Nor and Zagan Nor in central Mongolia (about long. 


100-102 E.). It has apparently disappeared from eastern Mon- 
golia (including adjacent parts of Siberia and Manchuria). Wild 
Asses throughout the world, with the apparent exception of the 
Tibetan Kiang, are a vanishing type. 

The following is derived from Pallas's early description (1781, 
pp. 16-17) , supplemented by Radde's description (1862, pp. 293-294) 
of what were virtually topotypical specimens from the vicinity of 
Dalai Nor: The general color of the summer pelage is reddish yellow, 
with a slight grayish tinge; in winter the color is more reddish than 
yellow, and the hair is longer. Snout whitish ; rest of head more and 
more yellowish ; mane brownish ; lower side of the neck of the general 
body color; upper rump ochraceous; limbs and ventral surface paler 
than sides; posterior side of forelegs, inner side of hind legs, lower 
rump, and posterior border of the thighs whitish ; a brownish-black 
median dorsal stripe from the mane to the bushy part of the tail, 
broadest on the hindquarters; bristly hairs above the hoofs blackish. 
Height at shoulder, about 3 feet 10 inches; length of ears, about 7 
inches; length of tail without hairy tuft, about 1 foot 4 inches. 

The former range apparently covered the greater part of Outer 
Mongolia (except the present Tannu-Tuva) , small areas in Siberia 
and Manchuria adjacent to the northeastern corner of Mongolia, at 
least the western part of Inner Mongolia, and the northern part of 
Chinese Turkestan (chiefly north of the Tian Shan) . 

According to Pallas (1781, pp. 5-8), the Argun steppes are the 
only place where these animals are still met with in Siberia. From 
the rest of Dauria, where they once ranged, they have retreated into 
the Mongolian deserts, on account of settlements. They still swarm 
about Tarei Nor. Formerly they were seen on the Argun steppes in 
great herds, but now only as solitary individuals or in scattered 
troops. On the Mongolian Gobi they occur in numerous herds. This 
is a game animal for the Mongols and Steppe Tungus, who eat the 
flesh and make boots of the hide. 

Radde writes (1862, p. 293) that in the fall and winter of 1856 a 
strong northward migration extended to the region between Tarei 
Nor and Dalai Nor, and that several animals were taken north of 
Dalai Nor- (in northwestern Manchuria) . In a rare journal (Beitrage 
Kenntniss Russ. Reiches, vol. 23, pp. 431-433) Radde gives addi- 
tional information on life history, hunting, and economic uses. 

A dearth of recent records of Wild Asses in eastern Mongolia 
bodes ill for their survival in that region. They are evidently gone 
from the adjacent parts of Siberia and Manchuria. Arthur de C. 
Sowerby (in Hit., March 14, 1938) believes they have ceased to 
exist in all these areas. 

In 1887, in the region about the southern base of the eastern Altai, 


at about longitude 96-100 E., Younghusband (1888, p. 495) saw 
"considerable numbers of wild asses." 

In the central Gobi, in 1922-25, R. C. Andrews (1924, pp. 152-154; 
1926, pp. 132-145, 299-302, 317-318) found considerable numbers 
of Wild Asses in the vicinity of Orok Nor and Zagan Nor (about 
long. 100-102 E.) "During the first two years of our work in the 
Gobi, we never saw wild asses in herds of more than fifteen or 
twenty, but we did not arrive in their country until after the breed- 
ing-season. In 1925 the herds numbered thousands. Evidently 
they collect at favorable localities just before the young are born . 
. . . The young are dropped about the beginning of July, and the 
asses seek a flat plain, undoubtedly for protection from wolves." 
(P. 302.) 

"I have been asked by many people if it would be possible to 
catch wild asses when they are young and use them for breeding 
purposes. I do not believe that this would be practicable, due to 
the extraordinary wildness of the animals. Certainly, it would be 
difficult to tame an adult wild ass." (R. C. Andrews, 1924, p. 154.) 

In 1926 the Kulan was very common at the northern base of Iche- 
Bogdo, in the valley of the lakes west of Orok Nor, and in the desert 
area to the northwest as far as the Baidarik River (Formozow, 
1931, p. 77). 

In 1911 Carruthers (1913, p. 532) found that the western shore 
of Bar Kul, in southern Dzungaria, was the haunt of droves of 
Wild Asses. His companion, J. H. Miller, supplies much additional 
information (in Carruthers, 1913, pp. 582, 588-589, 603-608). In 
the hills west of Bar Kul, towards the end of April, "a few wild- 
asses, straight from their winter quarters on the lowlands to the 
north, were busy making up for their scanty winter fare" (p. 588) . 
In the vicinity of Shi-Kho, at the northern base of the Tian Shan, 
a domesticated Kulon was examined; it was perfectly docile, but 
could not be broken to the saddle. A large wild herd was seen in the 
same area. (P. 603.) In Guchen a Kirghiz reported Kulon very 
numerous in the sand-dune area to the north, and Miller himself 
found fair numbers there (p. 604) . Two specimens were secured near 
the Dzungarian Gate north of Ebi Nor, where a spring was much fre- 
quented by Kulon (p. 605) . 

"My specimens are undoubtedly Equus hemionus typicus .... 
Its extreme eastern distribution is at present imperfectly known; 
Sir Francis Younghusband, in his journey across the Northern Gobi, 
mentions seeing kulon in the Gobi at the extreme eastern end of the 
Altai. They are found north of the Altai Range on the plains, 
round the large lakes in the Kobdo region; we met with them near 
Barkul, and in several other places throughout Southern Dzungaria. 
. . . The natives hunt them occasionally for their skins and meat, 


which they consider more palatable that the best mutton." (J. H. 
Miller, in Carruthers, 1913, pp. 607-608.) 

"The wild ass, or kulon, is unlikely to be seen unless a special 
attempt is made. . . . The kulon is a rare animal, excessively wild 
and lives in very difficult country. Featureless plains, bitterly cold 
in winter, waterless and sunbaked in summer, are its habitat. The 
kulon ranges . . . through Dzungaria to the edge of the Gobi. We 
have seen them at the lowest elevation in the heart of the conti- 
nent, and at 7,000 to 8,000 feet above the sea, in localities not very 
far distant from each other." (Carruthers, 1915, p. 154.) 

In 1926 a journey made by Lattimore (1929, pp. 228-321) 
through the southwestern Gobi filled in some blank spaces in the 
known distribution of this species. West of Edsin Gol, at the Wild 
Horse Well (about lat. 42 N., long. 99 E.) : "They say that on 
this fringe of the Khara Gobi there are wild horses . . . and wild 
asses" (p. 228). Near the "House of the False Lama" (about lat, 
42 30' N., long. 98 E.) : "To this whole series of springs there 
come at night antelope, wild asses, and, they say, wild camels" (p. 
243) . In the vicinity of Ming Shui (about lat. 43 N., long. 96 E.) : 
"To our camp that day there came riding a Mongol, who had fol- 
lowed us for two marches to sell the hinder half of a wild ass that 
he had shot in the Mongol way from a pit near the drinking place" 
(p. 251) . West of Ming Shui, near the eastern outposts of the Kar- 
lik Tagh: "Here the camel herders in the dawn reported a herd 
of wild asses. It was the only sight of them that I ever had .... 
Their skins make first-class clothing, with much more wear than the 
antelope skin. I have heard that there is a Turki proverb that wild 
asses are so hard to kill that even when you get the skin of one 
safely spread out on your sleeping platform it wiggles. The meat 
is something like beef, but a sublime beef. It is very dry, with a 
coarse grain and a strange aromatic sweetness. Chinese and Mon- 
gols put it above any other game, and it undoubtedly ranks with 
the noblest vension." (P. 252.) At Wu-t'ung Wo-tze, about 100 
miles northeast of Kucheng, in the Dzungarian Gobi: "It . . . was 
formerly a well-known wild-ass ground; but the wild ass in this 
region has been almost killed off by the Qazaqs. Both Mongols and 
Qazaqs will put themselves to more trouble to bag wild ass than 
almost any other game." (P. 321.) 

Farther southwest than the territory covered by Lattimore, along 
the route from Hami to Bulundsir River, Wild Asses were 
reported in 1898 in a number of places by Futterer (1901, pp. 179, 
180, 184, 188) . A specimen obtained northwest of the last-mentioned 
locality became the type of Equus hemionus luteus Matschie. 
In 1934 Sven Hedin (1940, pp. 195, 197, 200) found tracks in the 
Ghashun Gobi about 75 miles west of Futterer 's route. This area 


seems to constitute the southwestern limit (as far as known at 
present) of the range of the Mongolian Wild Ass. 

Enemies. Among predatory animals, the Wolf seems to be the 
only enemy of any importance, and doubtless it has never affected 
the Wild Asses at all seriously. Apparently it cannot successfully 
attack any except the young Asses within a few weeks of their 
birth. Older animals are able to outrun the Wolf on the open plains. 

Increasing use and precision of firearms in the hands of the 
Asiatics have undoubtedly contributed chiefly to the decline of the 
AVild Asses. 

Transcaspian Wild Ass; Transcaspian Kulan; Wild Ass of 
Russian Turkestan. Transkaspischer Kulan (Ger.) 


Equus (Asinus) hemionus finschi Matschie, in Futterer, Durch Asien, 
vol. 3, pt. 5, Zoologie (Nachtrag), p. 24, 1911. ("Nordostlich vom Saisan- 
nor" (Zaisan Nor, in former Province of Semipalatinsk, Russian Turke- 
stan).) (Cf. Harper, 1940, p. 198.) 

FIGS.: Radde, Sammlungen Kaukas. Mus., vol. 1, Zoologie, pi. facing p. 60, 
1899; Brehm's Tierleben (IV), 12, p. 670, tab. Unpaarhufer V, fig. 2, 
1915; Schwarz, 1929, p. 92, fig. 5. 

This Wild Ass is now very scarce in Russian Turkestan, having 
evidently disappeared from the greater part of the country. It was 
long ago exterminated in southern Russia. 

Matschie describes the type (from the vicinity of Zaisan Nor) 
as reddish salmon, with a slight tinge of gray; the lips are white; 
the whitish of the under parts extends well up on the flanks; the 
dark vertebral stripe continues on to the base of the tail. Schwarz 
adds (1929, p. 91) that the maximum width of the vertebral stripe 
is 42 mm. Radde and Walter (1889, p. 1059) describe a full-grown 
male from the Askabad region as lacking a shoulder stripe; its 
height at the shoulder was 1,110 mm.; tail (including tuft) , 590 mm. 

"It is clear [from Strabo's account] that the wild ass (onager) 
existed all across southern Russia in the fifth century B. C., for it 
was hunted both by the Sarmatian tribes who lived on the east side 
of the Don (Tanais) and by the Scythians who occupied the region 
to the west of that river. It is even possible that the wild ass dwelt 
in the Danube valley almost down to the beginning of the historical 
period. It seems certain that neither Sarmatian nor Scythian ever 
domesticated the wild ass, a circumstance probably due to the fact 
that they had a more docile and serviceable animal in the wild 
horses of the same region." (Ridgeway, 1905, pp. 51-52.) 

"In former days kulan and onagers appear to have ranged much 
further westward than is the case at the present day. It is stated, 
for instance, by the Russian naturalist Rytschkov that in the